LIBRARY 

OF  THE 

UNIVERSITY  OF  CALIFORNIA. 

GIFT    OF 


Class   3 


TERATOLOGY  IN  THE   FLOWERS  OF  TWO 
CALIFORNIAN  WILLOWS 


BY 

WILLIAM  WARNER  MOTT 


THESIS    PRESENTED    TO    THE    FACULTY    OF    THE    COLLEGE   OF    NATURAL 
SCIENCES     IN     PARTIAL     FULFILLMENT    OF     THE     REQUIRE- 
MENTS FOR  THE  DEGREE  OF  MASTER  OF  SCIENCE. 


UNIVERSITY   OF   CALIFORNIA    PUBLICATIONS 
BOTANY 

Vol.  2,  No.  7,  pp.  181-226,  Pis.  19-20  December  16,  1905 


TERATOLOGY  IN  THE   FLOWERS   OF  TWO 
CALIFORNIAN  WILLOWS 

BY 

WILLIAM  WARNER  MOTT 


CONTENTS. 

PAGE 

I.     Introduction 181 

II.     A  Teratological,  mainly  Staminate,  Salix  lasiolepis  Benth 184 

III.  A    Teratological,    mainly    Pistillate,    Salix    lasiandra    Benth.  X 

~babylonica  L.  and  some  related  individuals 196 

IV.  Bearing  of  the  Abnormalities  described  on  some  Moot  Points 

regarding  the  Salix  Flower 203 

V.     Some  Possible  Causes  of  the  Abnormalities  described 200 

VI.     Summary 213 

VII.     Bibliography 216 

VIII.     Explanation  of  Plates    219 


Introduction. 

The  normal  flowers  of  the  numerous  species  of  willows  are 
unisexual,  and  the  staminate  and  the  pistillate  flowers  are  born 
on  separate  individuals  (species  dioecious).  There  is  no  peri- 
anth, but  the  flowers  are  born  in  catkins,  and  the  bracts  of  a 
catkin  closely  subtend  the  sex-organs  composing  the  flowers  and 
protect  these  organs  in  the  young  stage.  The  number  of 
stamens  in  a  flower  varies  in  different  species,  but  two,  three, 
or  five  are  common.  The  pistil  is  unicellular  and  bicarpellary 
with  two  parietal  placentae,  and  the  fruit  is  a  capsule.  A  small 
honey-gland  is  borne  at  the  base  of  the  pistil  and  of  the  stamens, 
on  the  side  next  to  the  axis  of  the  catkin,  and  in  some  species  a 
second  basal  gland  is  borne  on  the  bract  opposite  to  the  first. 


172945 


182  University  of  California  Publications.  [BOTANY 

Cases  of  monoecism  (male  and  female  flowers  on  one  indi- 
vidual), and  of  abnormal  bisexual  flowers  with  perfectly  de- 
veloped pistil  and  stamens,  and  of  flowers  with  monstrous  sex- 
organs  combining  the  characters  of  pistil  and  stamens  in  vari- 
ous proportions  have  been  frequently  recorded  in  this  genus 
(Salix)  and  several  writers  have  speculated  at  some  length  as 
to  the  cause,  and  as  to  the  significance,  morphological,  anatomical, 
and  physiological,  of  these  abnormalities.  A  few  of  the  articles 
published  have  been  accompanied  by  illustrations,  notably  a 
paper  by  Camus  (1899)  and  two  by  von  Seemen  (1886,  1895), 
but  none  of  the  drawings  show  in  detail  a  series  of  the  monstrous 
organs  connecting  stamens  and  pistil.  The  object  of  this  paper 
is  mainly  to  present  drawings  of  such  a  series  with  particular 
note  of  certain  facts  well  shown  by  the  material  at  hand,  some  of 
which  have  not  been  found  described  elsewhere.  The  material 
was  discovered  by  the  writer  on  a  predominantly  male  individual 
of  Salix  lasiolepis  Benth.  (near  Berkeley,  in  the  spring  of 
1903).  The  male  tree  of  this  species  has  normally  two  stamens 
to  a  flower,  and  this  abnormal  specimen  showed*  the  two  stamens 
in  various  stages  of  transformation  (so  it  appeared)  into  pistil. 
The  converse  case  of  apparent  transformation,  namely,  pistil 
more  or  less  changed  into  stamens,  has  also  been  found  by  the 
writer  on  a  predominantly  pistillate  willow-tree,f  which  is  be- 
lieved to  be  a  hybrid  of  Salix  lasiandra  Benth.  and  Salix  baby- 
lonica  L. ;  and  on  this  tree  occurred  also  bisexual  flowers  with 
perfectly  developed  pistil  and  stamens  (the  number  of  stamens 
being  commonly  two),  and  also  two  extra,  lateral  floral  glands. 
Some  notes  of  these  abnormalities  are  made. 

This  pistillody  of  the  stamens  and  staminody  of  the  pistil, 
and  the  occurrence  of  bisexual  flowers  with  normally  developed 
sex-organs  have  not,  it  is  believed,  been  hitherto  reported  in 
Salix  for  this  Coast,  nor,  therefore,  for  this  species  and  this 
hybrid.  Also  no  mention  has  been  found  for  this  genus  of  the 
intimate  association,  which  is  shown  here  in  the  Salix  lasiolepis 

*  The  tree  was  destroyed  soon  after  it  was  found,  by  the  opening  of  a 
street. 

t  The  tree  is  still  growing,  near  Berkeley.  It  was  found  the  same 
season  as  the  8.  lasiolepis  first  mentioned. 


VOL.  2]          Mott.— Teratology  in  Calif  ornian  Willows.  183 

series,  of  the  micro-  and  macrosporangial  tissues, — ovules  pro- 
truding through  pollen-sacs,  and  causing  malformations  in  their 
walls.*  Likewise  there  seems  to  be  no  record  for  Salix  of  the 
occurrence  of  four  floral  glands  in  a  flower. 

The  possible  bearing  of  the  irregularities  described  upoti 
several  matters  of  discussion  regarding  the  genus  Salix  will  be 
briefly  indicated, — such  questions,  for  example,  as  whether  the 
ancestral  flower  was  perfect,f  instead  of  unisexual,  or  whether 
even  complete ;  t  what,  if  the  flower  were  perfect,  was  the  num- 
ber of  stamens ;  and  whether  the  two  carpels  of  the  now  normal 
pistil  stand  median  §  or  transverse.  || 

The  facts  described  seem  of  interest,  too,  in  connection  with 
a  wider  biological  question  of  what,  in  general,  causes  to  be  pro- 
duced, from  the  apparently  common  undifferentiated  tissue,  in 
one  case,  pistil,  in  another,  stamen,  in  another,  leaf,  stem,  or 
root, — the  question,  that  is,  of  what  are  the  determining  factors 
in  the  development  of  this  embryonic  basal  tissue  which  seems 
to  possess  the  potentiality  of  producing  all  forms  of  organ-tissue 
peculiar  to  a  species. 

The  occurrence  of  bisexual  flowers  with  distinct  pistil  and 
stamens,  on  these  willows,  in  place  of  normal  unisexual  flowers, 
and  the  abnormal  development  of  ovules,  female  bodies,  in- 
stead of  pollen-sacs,  male  bodies,  and  also  of  both  ovules  and 
pollen-sacs  at  once,  from  the  embryonic  tissue  of  the  connective', 
are  matters  of  interest  in  connection  with  the  general  problems 
of  sex, — not  only  in  the  plant  world  but  in  the  animal  world  as 
well.  In  the  animal  world,  also,  cases  of  casual  and  of  normal 
hermaphroditism,  of  different  degrees  of  intimacy,  occur. 

The  perfect  flowers  with  distinct  pistil  and  stamens  suggest, 
among  animals,  the  case  of  one  of  the  toads,  the  male  of  which 

*  Two  cases  somewhat  similar,  of  polleniferous  ovules  in  passion-vines 
(Passiflora  coerulea  and  P.  palmata)  have  been  described  and  illustrated 
by  Salter  (Trans.  Linn.  Soc.  XXIV,  p.  143,  tab.  XXIV),  and  a  case  of  the 
same  monstrosity  in  a  rose  (Kosa  arvensis)  has  been  reported  by  Masters 
(Seemann's  Journal  of  Botany,  1867,  p.  319,  tab.  LXXII,  figs.  B  1-9). 
An  instance  of  ovuliferous  anthers  in  a  gourd  (Cucurbita)  is  mentioned 
by  Berkeley  (Gard.  Chron.  1851,  p.  499.) 

t  Perfect :  having  the  essential  organs,  viz.  pistil  and  stamens. 

+  Complete:  having  pistil,  stamens,  and  also  perianth. 

§  Median:  in  the  line  of  the  bract  and  the  axis  of  the  ament. 

!l  Transverse:  in  a  line  at  right  angles  to  the  median  line. 


184  University  of  California  Publications.  [BOTANY 

has  frequently  a  rudimentary  ovary  in  front  of  the  testes,  and 
again  the  case  of  the  leech,  which  has  regularly  both  ovaries 
and  testes  in  one  individual,  the  two  sets  of  organs  being  quite 
distinct  from  each  other.  Many  fishes,  such  as  the  cod,  and.  a 
little  lower,  the  hagfish  are  often  similarly  hermaphrodite. 

The  case  of  the  monstrous  bisexual  floral  organs,  where  pollen- 
sacs  and  ovules  develop  with  apparent  indifference  from  con- 
nective tissue,  is  similar,  in  some  measure,  to  the  case  of  the 
regularly  hermaphrodite  snail,  where  sperm  and  ova  are  derived 
from  identical  cells  originating  in  the  so-called  hermaphroditic 
gland,  the  sexual  differentiation  of  these  reproductive  cells  de- 
pending merely  upon  which  of  two  glands  they  are  conducted 
into  for  their  development. 

In  degree  of  anatomical  intimacy,  the  hermaphroditism  seen 
in  the  monstrous  floral  organs, — where  the  seats  of  pollen-sac- 
and  ovule-production  are  so  very  close  together  on  one  phyllome, 
— is  analogous  to  that  bisexualism  occasionally  found  in  the 
common  frog,  where  a  testis  may  have  an  anterior  ovarian  por- 
tion, or  an  ovary  be  anteriorly  replaced  by  a  testis. 

These  cases,  then,  of  sexual  abnormality  among  plants,  like 
the  similar  instances  among  animals,  bear  upon  various  general 
questions  of  sex: — is  hermaphroditism  the  primitive  condition 
in  all  organisms ;  and  are  male  and  female  sex-elements, — micro- 
spores  and  macrospores,  sperm  and  ova, — universally  devel- 
oped by  differentiation  of  sexless  cells,  or,  better  perhaps,  of 
cells  potentially  of  either  sex ;  and  so  on.  The  facts  here  noted 
may  prove  of  interest  in  some  such  connection. 

A  Teratological,  Mainly  Staminate,  Salix  lasiolepis  Benth. 

PlSTILLODY  OP  THE  STAMENS  : DESCRIPTION  OP  A  SERIES  OP  SEX- 
UAL ORGANS  CONNECTING  THE  TWO  STAMENS  (THE  TYPICAL  MALE 
FLOWER  OP  THE  SPECIES )  WITH  A  BICARPELLARY  PISTIL  (THE  TYPI- 
CAL FEMALE  FLOWER  OF  THE  SPECIES ). — This  series  of  forms 
showing  increasing  amounts  of  pistillody  of  stamens  is  repre- 
sented in  the  series  of  figures  on  plates  19  and  20  (figs.  1-15). 
Only  enough  of  the  abnormal  forms  are  shown  to  make  evident 


VOL.  2]          Mott.— Teratology  in  Calif ornian  Willows.  1.85 

the  connection  between  stamens  and  pistil;  numberless  other 
phases  might  have  been  interpolated.  Figures  1  and  15  are  from 
normal  trees,  as  types  of  the  male  and  female  flowers,  for  com- 
parison. The  various  forms  that  occur  in  this  abnormal  tree, 
it  is  well  perhaps  to  state  clearly  at  once,  are  found  composing 
as  many  distinct  flowers.  That  is,  a  given  pair  of  sexual  phyl- 
lomes  do  not  go  through  a  transformation  from  a  perfectly  devel- 
oped pair  of  stamens,  to  a  perfectly  developed  pair  of  carpels 
forming  a  pistil,  but  remain  constantly  of  the  general  character 
that  marks  them  in  the  floral  bud,  *  until  they  finally  fall  with 
the  catkin.  However,  for  the  sake  of  vividness  in  bringing  out 
certain  points  that  are  shown  by  the  series  of  forms,  the  differ- 
ent members  of  the  series  are  described  as  though  each  repre- 
sented a  temporary  stage  in  a  progressive  change  from  stamens 
to  pistil  taking  place  in  a  single  flower.  To  follow  a  description 
of  such  a  process  of  change  it  is  well  perhaps  first  to  have  more 
clearly  in  mind  the  initial  and  final  stages. 

The  normal  staminate  flower  in  this  species  (cf.  pi.  19,  fig.  1) 
consists  of  the  two  stamens,  standing  transverse,  with  cylindrical 
filaments  more  or  less  united.  The  amount  of  union  of  the  fila- 
ments is  variable  in  the  species.  In  the  flower  chosen  to  show 
the  normal  type,  the  filaments  are  united  about  to  the  middle 
as  in  the  case  of  the  most  nearly  normal  staminate  flowers  on 
our  abnormal  tree.  The  anthers  are  adnate, — that  is,  attached 
by  their  whole  length  to  the  ends  of  the  filaments.  Each  anther 
is  composed  of  four  pollen-sacs  (or  microsporangia),  in  two 
pairs,  the  two  pollen-sacs  of  a  pair  being  closely  united  to  each 
other,  and  shedding  pollen  through  a  single  slit.  The  anthers 
in  the  young  stage  are  back  to  back  (extrorse)  (cf.  fig.  IA,  pi. 
19),  but  by  the  time  they  are  ready  to  shed  their  pollen,  they 
have  turned,  through  torsion  of  the  filaments,  to  face  the  honey- 
gland. 

The  normal  female  flower  (cf.  figs.  14  and  15)  consists  of 
the  unicellular,  bicarpellary  pistil.  The  two  carpels  stand — it 

*  Whether  a  pair  of  phyllomes  which  are  finally,  at  least,  abnormal 
undergo  such  a  change,  more  or  less  completely,  in  the  very  young  stage  in 
the  floral  bud,  or  whether  the  pair  are  abnormal  from  their  very  inception 
at  the  meristem — these  are  points  apparently  still  to  be  worked  out. 


186  University  of  California  Publications.  [BOTANY 

is  commonly  considered — transverse,  like  the  stamens.  The  stalk. 
or  stipe,  is  short.  The  two  placentae  are  median,  and  on  the 
walls  of  the  ovary  (parietal),  and  only  on  the  lower  part  of 
the  walls  (see  fig.  13).  The  number  of  ovules  is  from  six  to 
eight.  The  upper  barren  portion  of  the  ovary  is  tapering-,  and 
affords  space  for  the  tufts  of  hair  arising  from  the  ovule  stalks, 
or  funiculi.  The  style  is  single  and  rather  short,  and  the  stigrma 
is  more  or  less  bilobed  transversely.  Two  longitudinal  seams, 
one  in  the  anterior  *  wall  of  the  ovary  and  one  in  the  posterior  f 
wall,  mark  the  sutures  between  the  two  carpels  and,  as  well,  the 
position  of  the  two  placentae  within.  Two  slight  longitudinal 
grooves,  on  the  right  and  left  sides  of  the  ovary,  follow  the 
midribs  of  the  carpels.  It  is  along  these  lateral  grooves,  and 
not  at  the  sutures,  that  dehiscence  occurs  (dehiscence  loculicidal, 
see  fig.  12)  ;  hence  the  two  valves  of  the  capsule  are  median  and 
the  seeds  are  found  on  the  middle  of  the  valves. 

The  change  from  stamens  to  pistil  is  effected,  in  brief  (a 
detailed  description  follows  later),  by  a  median  broadening  and 
also  an  apical  development,  and  final  union,  of  the  two  con- 
nectives, to  form  the  ovary  of  the  pistil — (1)  the  lower  ovule- 
bearing  portion  and  (2)  the  upper  barren  portion — and  also 
(3)  the  style  and  (4)  the  stigma,  and  further  by  a  shortening, 
and  completion  of  the  fusion,  of  the  filaments,  to  form  the  stipe 
of  the  pistil.  The  vegetative  broadening  of  the  connectives 
occurs  in  such  a  way  as,  at  an  early  stage  in  the  series,  to  throw 
the  reproductive  portions  inward,  so  that  the  anthers,  which  are 
produced  from  these  portions,  are,  in  the  young  stage,  face  to 
face  (introrse)  (though  normally  extrorse)  and  the  ovules,  pro- 
duced from  the  same  reproductive  regions,  are,  finally,  enclosed 
when  the  connectives  unite.  The  change  in  the  general  form 
of  the  phyllomes  from  that  of  a  stamen  to  that  of  a  carpel  is 
proportional  (with  some  latitude)  to  the  degeneration  of  the 
pollen-sacs,  and  the  replacement  of  them  by  ovules.  That  is, 
all  the  male  characters  of  the  phyllome,  being  correlated,  vary 
approximately  together,  and  the  female  characters  the  same. 


*  Anterior :  toward  the  bract. 

t  Posterior :  toward  the  axis  of  the  catkin. 


VOL.  2]          Mott— Teratology  in  Calif ornian  Willows.  187 

The  two  phyllomes  of  a  flower  usually  show  the  same  amount 
of  change,  but  they  may  differ  somewhat.  Dehiscence  of  the 
ovary,  so  long  as  it  is  monstrous  to  the  extent  of  bearing  anthers, 
or  anther-vestiges,  is  premature  and  at  the  sutures,  so  that  the 
transversely  standing  phyllomes  remain  intact ;  after  the  anthers 
have  quite  disappeared,  the  dehiscence  is  no  longer  premature, 
and  it  occurs  at  the  midribs  (as  in  the  flower  on  the  normal  tree) 
so  that  the  two  valves  of  the  capsule  are  anterior  and  posterior. 

A  detailed  account  of  this  change  from  stamens  to  pistil  is 
best  given  by  describing  in  order  the  forms  represented  in  the 
series  of  drawings.  Various  facts  that  are  exhibited  by  the 
series  as  a  whole,  or  by  certain  parts  of  it,  will  be  touched  upon 
under  those  figures  that  are  most  suggestive  of  the  particular 
facts.  In  all  the  figures,  it  is  the  posterior  side  of  the  flower 
which  is  shown, — the  side  toward  the  axis  of  the  ament.  The 
bract,  honey-gland,  and  lower  part  of  the  united  filaments  are 
in  most  cases  not  shown.  Most  of  the  figures  are  about  ten  times 
natural  size. 

Fig.  1.  This  is  a  typical  staminate  flower  from  a  normal 
tree.  It  has  already  been  described.  The  four  pollen-sacs  of 
each  anther  are  seen  to  be  closely  associated.  Later  on  in  tlie 
series,  these  pollen-sacs,  or  microsporangia,  degenerate,  and  the 
vestiges  often  stand  independent  of  each  other,  as  their  proto- 
types in  the  Gymnosperms  and  Cryptogams  quite  commonly  do. 
Fig.  IA  shows  a  young  diandrous  flower  (S.  purpurea  L.)  with 
extrorse  anthers. 

Fig.  2.  This  is  the  nearest  approach  to  absolutely  normal 
flower  that  is  found  on  the  tree.  It  is  a  very  common  type.  The 
free  ends  of  the  two  filaments  (which  we  will  call  the  "forks") 
are  apparently  never  perfectly  cylindrical  but  are  always  slightly 
grooved  longitudinally  on  the  sides  that  face  each  other.  These 
two  grooves  are  the  first  indications  of  the  formation  of  the 
ovarian  cavity  of  the  pistil.  As  has  been  said,  however,  it  is 
the  connectives  which  form  the  ovary,  and  later  in  the  series, 
these  forks  cease  almost  entirely  to  be  developed,  the  pair  of 
filaments  as  a  whole  shortening  to  become  the  stipe  of  the  pistil. 
These  two  grooves  at  this  stage  arise  as  follows : — 


188  University  of  California  Publications.  [BOTANY 

While  in  the  floral  bud  (and  these  first  remarks  are  true 
of  stamens  in  general)  the  filaments  attain  about  their  full  de- 
velopment in  thickness,  but  remain  very  short,  until  the  anthers 
have  reached  almost  their  full  size  and  are  nearly  ready  to  dis- 
charge their  pollen.  This  keeps  the  anthers  within  the  protec- 
tion of  the  bract  as  long  as  possible.  Just  before  dehiscence 
of  the  pollen-sacs,  the  cells  of  the  filaments  increase  greatly  in 
length,  so  that  the  anthers  are  exserted  and  the  pollen  is  ren- 
dered accessible  to  insects.  Now  while  these  abnormal  stamens 
are  still  in  the  bud,  and  the  filament  forks  are  short  (remem- 
bering that  the  anthers  of  the  abnormal  phyllomes,  at  least 
while  young,  are  introrse  as  explained  later)  there  is  a  hollow 
on  the  inner  side  of  each  fork,  at  the  base  of  the  anther.  The 
presence  of  this  hollow  is  due  to  the  fact  that  the  connectives, 
which  bear  the  anthers  directly,  have  commenced  to  become 
grooved  on  the  inner  sides  preparatory  to  the  formation  of  the 
concave  ovary  walls,  and  this  grooving  has  extended  downward 
into  the  short  filament  forks.  Hence  as  the  cells  of  the  forks 
lengthen,  these  slight  hollows  develop  as  grooves  along  the  whole 
length  of  the  forks. 

The  anthers  here  have  twisted  to  face  the  honey-gland,  and 
the  flower  is  otherwise  normal. 

Fig.  3.  The  forks  and  connectives  here  are  broadened  some- 
what, medianly,  and  the  grooving,  or  concaving,  is  more  pro- 
nounced. The  anthers,  which  are  still,  from  the  bud.  largely 
facing  each  other,  are  seen  to  be  introrse  in  their  position  on 
the  connectives;  that  is,  the  four  reproductive  portions  of  each 
broadening  connective,  and  hence  the  four  pollen-sacs,  have  been 
brought  to  face  inward.  This  change  is  anticipatory  of  the 
production  of  ovules — which  are  to  come  from  these  same  repro- 
ductive portions — for  the  ovules  are  finally  to  be  enclosed  by  the 
union  of  the  broadened  connectives.* 


*  The  fact  that  the  anthers  in  these  abnormal  forms  are  introrse,  at 
least  in  the  young  stage,  instead  of  extrorse  as  in  the  normal  flower  of  the 
species,  it  is  possible,  it  seems,  to  account  for  as  follows: 

In  the  young  stamens — and  this  is  true  of  stamens  in  general  of  this 
simple  Salix  type — the  four  pollen  sacs,  or  microsporangia,  are  produced 
by  a  differentiation  longitudinally  of  the  tissue  at  the  end  of  the  stamen. 
Two  pollen  sacs  are  formed  in  the  anterior  angles  of  the  stamen,  one  right 


VOL.  2]          Mott.— Teratology  in  Calif ornian  Willows.  189 

The  anthers  are  seen  to  have  commenced  to  twist,  mainly 
through  mere  torsion  of  the  forks,  to  face  toward  the  honey- 
gland.  The  tendency  of  the  anthers  to  assume  this  position, 
when  they  are  mature,  is  present  in  the  normal  flower,  and  is 
shown  all  through  the  series  of  abnormal  forms  as  long  as  any 
trace  of  the  pollen-sacs  remains.  The  advantage  of  the  position 
for  insect  pollination  is  obvious. 

The  posterior  margins  of  the  forks  are  ununited.  Either  the 
twisting  of  the  anthers  to  face  the  honey-gland  has  torn  them 
apart,  or  else  the  swelling  of  the  still  well-developed  anthers 
prevented  them  from  ever  uniting.  This  condition,  namely,  of 
intact  anterior  suture  and  prematurely  ruptured,  or  never 
formed,  posterior  suture, — is  common  in  these  monstrous  forms 
especially  later  in  the  series.  In  the  earlier  forms  of  the  series, 
while  the  anthers  are  still  well  developed,  there  is  usually  no 
union  of  the  forks  at  all. 

Fig.  4.  This  figure  shows  the  first  appearance  of  ovules, 
and  the  ovules  are  seen  to  be  close  to  the  basal  end  of  the  anther- 
lobes.  Only  one  of  the  stamens  bears  them.  The  two  phyllomes 
of  a  flower  are  not  infrequently  different  in  character  to  the 
extent  shown  here.  The  right  stamen  is  similar  to  both  phyl- 
lomes in  fig.  2,  though  the  groove  is  a  little  more  evident.  The 
left  fork  has  reached  a  little  further  developed  stage.  The 
ovules  are  seen  to  point  inward.  There  is  only  one  ovule,  as  yet, 
on  each  margin  of  the  phyllome.  The  normal  number  on  each 

and  one  left,  and  two  in  the  posterior  angles,  one  right  and  one  left,  the 
portion  of  the  stamen  between  the  two  pairs  being  the  "connective."  The 
position  of  the  pollen-sacs  in  the  mature  state  of  a  stamen  is  often  dif- 
ferent from  this,  owing  to  the  later  vegetative  processes  of  growth  in  the 
stamen-tissue,  especially  in  the  region  of  the  connective.  These  are  well- 
established  facts.  Now  in  the  normal  flower  of  this  diandrous  willow,  the 
two  stamens  are  close  together,  and  it  is  apparently  because  the  pollen-sacs 
crowd  against  each  other  as  they  develop  and  swell  with  pollen,  that  they 
come  to  face  outward,  or  become  extrorse.  In  the  bud,  the  two  extrorse 
anthers,  standing  close  together,  appear  as  in  fig.  IA,  pi.  19.  In  the 
series  of  abnormal  forms,  where  the  two  connectives  gradually  become 
broadened  medianly  to  form  the  concave  ovary-wall,  it  is  apparently  be- 
cause the  vegetative  increase  necessary  to  effect  this  broadening  is  greatest 
on  that  side  of  each  connective  which  is  farthest  from  the  other  connective, 
that  the  four  pollen-sacs  are  thrown  to  face  inward,  or  become  introrse. 

The  connectives  in  fig.  3  are  not  yet  greatly  broadened  but  there  seems 
to  have  been  sufficient  growth  on  their  outer  sides  to  throw  the  pollen-sacs 
inward. 


190  University  of  California  Publications.  [BOTANY 

margin  of  a  carpel  is  three  or  four,  since  the  normal  numbor 
on  each  placenta  of  the  pistil  is  six  or  eight  and  the  placenta 
is  formed  by  the  union  of  two  carpel  margins.  That  these  two 
bodies  are  undoubtedly  ovules  is  shown  by  the  enlarged  drawing, 
fig.  16.  This  drawing  represents  an  optical,  longitudinal,  me- 
dian section  of  one  such  body  through  the  raphe.  A  similar 
section  of  an  ovule  from  a  pistil  borne  on  a  normal  tree  is  iden- 
tical with  this.  The  characters  are  unmistakable: — central  nu- 
cellus  enclosed  by  two  coats  (integuments),  basal  micropyle 
(ovule  anatropus),  and  raphe,  or  prolongation  of  the  stalk  (funi- 
culus),  with  its  vascular  bundle  terminating  at  the  (chalaza) 
base  of  the  two  coats. 

Fig.  5.  Here  the  number  of  ovules  has  increased,  and  the 
placental  vascular  strands  have  become  evident  as  thickenings 
on  the  margin  of  the  forks.  The  vascular  bundle  in  a  normal 
stamen  is  so  little  developed  as  scarcely  to  be  detected,  for  only 
enough  nutritive  material  has  to  be  conducted  to  the  pollen-sacs 
(microsporangia)  to  bring  them  to  maturity,  while  to  the  ovules 
(macrosporangia)  sufficient  food  has  to  be  conducted  not  only 
to  bring  them  to  maturity  as  macrosporangia,  but  also  to  supply 
their  demands,  after  their  fertilization,  by  the  pollen,  while  they 
are  being  developed  to  maturity  as  seeds. 

There  is  an  ovule  at  the  base  of  each  half -anther,  or  anther- 
lobe,  and  the  bodies  a  and  c  are  ovules  even  protruding  through 
the  anthers,  so  closely  are  the  ovules  associated  with  the  pollen- 
sacs  in  the  production  of  both  from  the  reproductive  portions 
of  the  phyllomes.  In  fact  it  is  this  close  association,  which  is 
seen  commonly  in  the  forms  of  this  type,  that  shows  that  it  is 
a  fairly  definite  part  of  a  phyllome  which  is  reproductive.  In 
other  words,  the  connective  region  of  the  stamen  and  the  pla- 
cental regions  of  a  carpel  seem  to  be  homologous. 

The  small  bodies  higher  up,  in  the  sinuses,  or  furrows,  in 
the  face  of  the  anthers,  where  two  pollen-sacs  unite  to  form  a 
half  anther, — bodies  which  look  like  ovules, — are  in  reality  lobes 
in  the  walls  of  the  pollen-sacs.  There  seems  to  be  no  question 
that  these  lobes  are  caused  by  abortive  ovules  beneath,  for  though 
in  most  cases  there  is  nothing  but  pollen  within  such  lobes,  in 


VOL.  2]          Mott. — Teratology  in  Calif ornian  Willoivs.  191 

some  instances  there  is  a  slight  protuberance  on  the  connective 
beneath;  and  an  ovule,  in  general,  originates  in  this  way.  The 
cases  such  as  seen  here  (noted  above)  of  ovules  actually  pro- 
truding through  pollen-sac  tissue,  are  further  evidence  that  the 
lobes  are  caused  by  incipient  ovules.  The  tissue  &,  for  example, 
around  the  base  of  the  protruding  ovules  a  is  unmistakably 
anther-tissue,  and  it  would  doubtless  have  constituted  a  lobe  only 
that  the  ovule  succeeded  in  keeping  from  being  overwhelmed, 
as  ovule  and  pollen-sac  developed  together;  that  is,  the  crowding 
of  the  anther-tissue  here  was  partially  withstood  by  the  devel- 
oping ovule,  while  in  other  cases  anther-tissue  gained  almost 
complete  supremacy,  and  yet  lobes  in  the  pollen-sac  walls  were 
caused  by  the  overwhelmed  ovules.* 

The  pollen-sac  lobe  d  looks  particularly  like  an  ovule ;  it 
stands  out  in  the  sinus  almost  independent  of  the  main  portion  j 
of  the  pollen-sac.  The  lobe  e  on  the  right  half  of  the  right 
stamen,  on  the  other  hand,  is  merely  a  slight  elevation  in  the 
surface  of  the  wall  of  its  pollen-sac.  Other  lobes  (also  marked  e) 
are  more  or  less  intermediate  in  form.  (The  means  of  identify- 
ing pollen-sac  tissue  are  seen  in  figs.  17-19,  and  the  matter  of 
this  identification  is  further  referred  to  under  fig.  7.) 

The  right  anther  lobe  has  dehisced. 

Fig.  6.  The  forks  and  connectives  in  this  figure  have  become 
considerably  broader,  and  here,  on  the  left  phyllome  at  c,  is  seen 
the  first  indication  of  apical  development  of  the  connective  to 
form  the  upper  barren  portion  of  the  ovary,  and  the  style  and 
stigma.  The  right  connective  is  not  yet  so  developed.  (The 
ovules  are  usually  produced  nearer  to  the  anthers  than  shown 
here. ) 

Fig.  7.  The  right  phyllome  here  is  still  predominantly  stam- 
ineal,  the  anther  being  well  developed,  and  the  fork  elongated. 
The  left  phyllome,  however,  is  about  equally  divided  in  char- 
acter between  stamen  and  carpel.  The  anther  has  been  reduced 
to  vestiges,  and  the  fork  accordingly  is  only  partially  developed, 

*  These  effects  of  pollen-sacs  and  ovules  upon  each  other,  where  they  are 
produced  together,  are  such  as  one  might  expect,  in  consideration  of  the 
processes  by  which  pollen-sacs  and  ovules,  in  general,  are  produced  by  dif- 
ferentiation of  embryonic  tissue  of  stamen  and  carpel  respectively. 


192  University  of  California  Publications.  [BOTANY 

though  the  united  part  of  the  two  phyllomes  is  still  of  the  normal 
length  in  the  stamen.  The  development  of  pollen-sacs  and  the 
elongation  of  the  cells  of  the  filament  are  correlated  male  char- 
acters of  the  phyllome,  and  the  development  and  the  elongation 
vary  in  amount  approximately  together.  The  lengths  of  the 
cells  of  the  two  forks  in  this  case  were  found  to  be  about  in- 
versely proportional  to  the  lengths  of  the  forks.  On  the  left 
phyllome,  ovules  have  almost  entirely  supplanted  the  right  lobe 
of  the  anther,  all  that  remains  of  the  lobe  being  the  vestige  d. 
At  the  same  time,  there  has  occurred  a  considerable  apical  devel- 
opment of  the  connective  toward  the  formation  of  the  upper 
pistil-parts  (barren  portion  of  the  ovary,  style,  and  stigma), 
this  being  a  female  character  of  the  phyllome  correlated  with 
the  production  of  ovules.  The  ovules  here  are  all  well  devel- 
oped. The  beginning  of  the  upper  barren  portion  of  the  ovary 
is  seen  at  7i',  the  style  at  li,  and  the  stigma  at  i.  The  styles 
and  stigmas  of  the  two  phyllomes  unite  later  on.  The  style  here, 
being  still  free,  has  shriveled  and  curled  downward. 

The  body  d  on  the  left  phyllome  might  be  taken  superficially 
for  an  ovule,  it  being  minute  and  otherwise  similar  to  an  ovule, 
and  being  borne  among  ovules,  but  it  proves  under  the  higher 
power  of  the  microscope  to  be  a  pollen-sac  vestige;  just  as  the 
still  more  ovule-like  body  d,  pi.  19,  fig.  5,  proved  to  be  of  pollen- 
sac  tissue.  Even  under  the  lower  power,  the  ovules  appear 
slightly  fluted  longitudinally  and  the  oblong  shape  indicates  the 
polarity  they  possess;  while  the  pollen-sac  lobes  and  even  the 
least  developed  pollen-sacs  have  a  smooth  surface  of  small  poly- 
gonal cells  such  as  normally  developed  pollen-sacs  have.  Pollen- 
sac  lobes  and  vestiges  also  often  contain  grains  of  pollen,  normal 
at  least  superficially  (see  pi.  20,  fig.  18),  (though  these  are  often 
lost  in  dissecting  out  the  body  for  examination).  Under  the 
higher  power,  d,  for  example,  shows  the  cellular  wall  structure 
which  is  characteristic  of  the  normally  developed  pollen-sac. 
This  peculiar  structure  is  illustrated  by  figs.  17  and  19.  The 
wall,  is  composed  of  an  outer  layer  of  thin-walled  cells  and  an 
inner  layer  of  so-called  fibrous  cells.  On  approaching  maturity, 
the  outer  layer  dries  more  rapidly  than  the  inner,  and  contracts 


VOL.  2]          Mott. — Teratology  in  Calif ornian  Willows.  193 

more  (as  does  the  drying  film  of  clay  over  coarser  material  on 
the  site  of  a  recent  pool),  and  the  rupturing  along  the  weakest 
line,  namely,  the  suture  between  two  pollen-sacs,  is  dehiscence. 
Hence  the  identity  of  ovules  and  of  pollen-sac  vestiges  and  lobes 
is  established. 

Fig.  8.  The  connectives  here  are  broadened  almost  to  the 
full  amount ;  the  anterior  margins  are  united ;  a  style  and  stigma 
have  developed  from  each  connective,  though  they  are  mal- 
formed and  the  styles  and  the  stigmas  are  still  ununited;  the 
old  filament  forks  are  both  reduced,  though  the  old  united  por- 
tion of  the  filaments  is  still  longer  than  a  stipe.  The  anthers 
at  this  stage  are  usually  more  degenerate.  There  is  about  a 
normal  number  of  ovules,  on  the  anterior  placenta.  The  ovarian 
cavity  is  well  developed,  though  the  posterior  margins  of  the 
phyllomes  (which  were  very  likely  united  in  a  younger  stage) 
have  been  torn  apart.  This  figure  illustrates  the  fact  that  the 
monstrous  ovary  dehisces  prematurely  and  at  the  sutures,  in- 
stead of  at  the  midribs. 

Fig.  8.  The  connectives  have  now  both  reached  full  carpel- 
width,  both  anterior  and  posterior  margins  have  united  below, 
though  the  upper  parts  are  still  free.  The  tapering  upper  por- 
tion of  the  ovary  is  not  yet  well  developed,  and  the  figure  shows 
(better  than  the  last)  a  rather  common  persistence  of  abnorm- 
ality of  style  and  stigma.  The  anthers  have  become  so  nearly 
eliminated  that  dehiscence  is  no  longer  premature,  and  yet  seeds 
are  not  matured,  the  organ  falling  early.  As  a  whole,  the  two 
phyllomes  have  come  to  suggest  pistil  rather  than  pair  of  stamens. 

Fig.  10  shows  close  approximation  to  normal  pistil.  The 
phyllomes  are  wholly  united,  the  ovary  is  tapering  above,  and 
style  is  single  and  there  is  clear  demarcation  of  the  style  from 
the  ovary.  The  lobing  of  the  stigma,  however,  is  still  right  and 
left,  corresponding  to  the  original  stamens,  and  not  yet  to  the 
valves.  The  stipe  is  still  abnormally  long.  (Even  though,  how- 
ever, the  lobes  of  the  stigma  in  the  normal  pistil  in  this  species 
commonly  correspond  to  the  valves,  dehiscence  does  not  extend 
through  the  style  and  stigma,  but  style  and  stigma  either  fall 
intact  from  the  ripe  capsule  or  else  break  away  from  one  valve 
at  the  time  of  dehiscence,  and  remain  attached  to  the  other.) 


?x 

(   UNIVERSITY  ] 

V  OF  / 


194  University  of  California  Publications.  [BOTANY 

Fig.  11  shows  the  same  specimen  as  fig.  10,  but  with  the 
nearer  (posterior)  half  of  the  ovary  removed,  to  show  that  the 
pollen-sacs  have  completely  disappeared,  and  that  the  ovules 
are  in  their  normal  position:  that  is,  they  are  median,  for  they 
are  borne  on  the  margins  of  transverse  phyllomes,  as  we  have 
seen  in  the  preceding  figures ;  and  they  are  on  the  lower  part  of 
the  ovary-wall,  for  the  old  filament  forks — below  the  reproduc- 
tive portion  of  the  phyllomes — have  been  reduced  and  at  the 
same  time  the  old  connectives — the  reproductive  portions  them- 
selves— have  been  developed  apically  to  form  the  upper  barren 
portion  of  the  ovary-wall. 

Fig.  12  shows  the  completion  of  the  change.  The  stipe  is  of 
normal  length,  the  ovary  has  swollen  with  seeds  and  down,  and 
has  matured  as  a  normal  capsule  dehiscing  at  the  carpel  mid- 
ribs so  that  the  valves  are  anterior  and  posterior.  Style  and 
stigma  have  fallen  away.  The  valves  separate  as  normally,  only 
half  or  two-third's  of  the  way  down. 

Fig.  13  presents  the  anterior  valve  of  the  capsule  in  fig.  12, 
pi.  20,  with  basal  remnants  of  the  other  valve,  to  show  the  char- 
acteristic position  of  the  placenta, — median  and  at  the  lower 
half  of  the  suture  between  the  phyllomes. 

Fig.  14,  showing  a  pistil  from  a  normal  tree,  has  already 
been  described.  It  exhibits  the  typical  characters  which  we 
have  seen  developing  through  the  series.  The  view  of  the  pistil 
is  posterior,  as  in  the  preceding  figures,  for  the  sake  of  com- 
parison. 

Fig.  15  shows  the  same  flower  as  in  the  last  figure,  but  turned 
somewhat  to  bring  into  view  one  of  the  slight  grooves  at  the 
carpel  midribs  along  which  normal  dehiscence  occurs,  as  con- 
trasted with  the  abnormal  dehiscence  at  the  sutures  in  the  mon- 
strous forms. 

There  is  thus  a  complete  series  of  forms  on  this  abnormal 
Salix  lasiolepis  connecting,  by  slight  gradations,  a  pair  of 
stamens  and  a  bicarpellary  pistil. 

MANNER  OF  OCCURENCE  OF  THE  ABNORMAL  FLOWERS  ON  THE 
CATKINS,  AND  EFFECT  OF  THEIR  PRESENCE  ON  THE  TIME  OF  FALLING 
OF  THE  CATKINS. — The  flowers  abnormal  to  the  tree  are,  of 


VOL.  2]          Mott. — Teratology  in  Calif  ornian  Willows.  195 

course,  both  those  with  any  form  of  monstrous  hermaphrodite 
organ  and  also  those  with  perfectly  developed  pistil,  for  the  tree 
as  a  whole  was  evidently  predominantly  staminate.  The  pre- 
dominance of  the  male  character  of  the  tree  as  a  whole  was 
shown  by  the  fact  that  when  the  tree  was  in  full  bloom,  anthers 
were  commonly  enough  present  in  the  flowers  to  give  almost  as 
much  of  a  yellow  cast  to  the  tree  as  is  characteristic  of  the 
normal  male  individual.  Many  of  the  catkins  were  composed 
wholly  of  practically  normal  male  flowers,  but  the  majority  of 
the  catkins  were  made  up  of  monstrous  hermaphrodite  flowers 
which  yet  were  more  staminal  in  character  than  pistillary. 
Quite  a  large  number  of  catkins  which  were  chiefly  of  monstrous 
flowers  showed  more  of  a  green  cast  than  yellow,  the  majority 
of  the  sex-organs  here  resembling  pistils  rather  than  pairs  of 
stamens.  Again  a  considerable  number  of  catkins  were  com- 
posed wholly  of  perfectly  developed  pistillate  flowers.  Then 
in  addition  to  these  four  sorts  of  catkins  named,  composed 
wholly  or  mainly  of  one  type  of  flower,  there  were  many  catkins 
showing  combinations  of  these  types  of  flowers  in  various  pro- 
portions; and  the  different  types  occuring  on  a  given  catkin 
might  be  rather  thoroughly  intermingled,  or  might  be  in  dif- 
ferent parts  of  the  catkin  (as  base,  middle,  and  tip),  but  without 
any  regularity  as  to  which  type  occurred  on  any  part. 

The  presence  of  monstrous  sex-organs  on  a  catkin  tended  to 
cause  the  catkin  to  drop  early, — earlier  even  than  a  wholly  male 
catkin  drops.  In  general,  of  course,  male  catkins  of  the  willow 
fall  earlier  in  the  season  than  the  female,  for  the  function  of 
the  staminate  catkins  in  fulfilled  earlier, — that  is,  as  soon  as  the 
matured  pollen  is  shed, — while  the  pistillate  catkins  must  per- 
sist, after  this  escaped  pollen  has  fallen  upon  the  stigmas,  until 
the  seeds  have  matured  and  scattered.  These  monstrous  her- 
maphrodite organs,  however,  unless  very  predominantly  of  one 
sex  or  the  other,  soon  shriveled  after  the  opening  of  the  flowers, 
so  that  catkins  wholly  of  monstrous  flowers  soon  fell  after 
blooming.  The  majority  of  the  catkins  borne  by  the  tree  being 
of  the  decidedly  monstrous  type,  the  tree  as  a  whole  had  few 
catkins  hanging  upon  it  at  a  time  (March  26)  when  normal  male 
individuals  near  bv  still  had  many  catkins  on  them  in  bloom. 


196  University  of  California  Publications.  [BOTANY 

A  peculiar  fact  noticed  was  that  a  very  few  normally  de- 
veloped pistils  on  a  catkin  were  sufficient  to  keep  the  catkin  from 
falling  until  the  maturation  of  the  seeds  in  these  pistils, — what- 
ever the  character  of  the  remaining  flowers,  male,  hermaphrodite, 
or  of  both  kinds.  Hence,  as  catkins  with  just  a  few  normal 
pistils  were  not  rare,  toward  the  end  of  the  season,  when  capsules 
on  trees  generally  were  dehiscing,  a  considerable  proportion  of 
what  catkins  still  remained  on  this  tree  displayed  only  very 
scattering  capsules ;  and  search  among  the  fallen  catkins  beneath 
the  tree  revealed  very  few  that  had  more  than  one  or  two  un- 
matured  pistils. 

Catkins  throughout  the  season,  whether  blooming  earlier  or 
later,  showed — in  about  the  same  proportion  of  cases — the  ab- 
normalities, arid  the  diversity  among  the  catkins  was  about  the 
same  throughout  the  season. 

A  Teratological,  Mainly  Pistillate,  Salix  lasiandra  Benth.X 
babylonica  L.*  and  Some  Related  Individuals. 

ON     THE     TREE     FIRST  *  NAMED  : STAMINODY     OF     THE     PISTIL ; 

BISEXUAL  FLOWERS  WITH  NORMALLY  FORMED  PISTIL  AND  STAMENS  ; 
EXTRA  FLORAL  GLANDS;  OTHER  NOTES;  THIS  TREE  AND  TWO  SIMI- 
LAR ONES  ARE  SCION-PROPAGATED. — Staminody  of  the  pistil  was 
one  of  the  abnormalities  shown  by  this  mainly  pistillate  tree. 
That  is,  in  the  normal  position  of  the  pistil  there  occurred 
monstrous  organs  which  possessed  in  various  proportions  the 
characters  both  of  pistil  and  of  stamens.  (See  a  pi.  20,  fig  21.) 
This  is  the  converse  of  the  apparent  transformation  of  sex- 
organs  seen  in  the  Salix  lasiolepis:  there  the  two  stamens  ap- 
peared more  or  less  changed  to  pistil,  here  the  pistil  is  more  or 
less  changed  to  two  stamens.  The  monstrous  pistils  vary  from 

*  There  are  (still  growing)  three  trees  of  this  general  description, 
standing  near  together  (at  the  corner  of  San  Pablo  Avenue  and  McAvoy 
Lane,  near  Berkeley),  which  were  grown  from  cuttings  planted  some 
fifteen  years  ago,  and  also  about  a  dozen  trees  of  similar  description,  but 
of  both  sexes,  standing  not  far  from  the  first  three  in  a  small  mixed  grove 
of  willows  (at  the  foot  of  Powder  Works  Hill),  which  were  naturally 
grown,  probably  from  seed.  The  three  cuttings  came  from  this  grove,  and 
were  probably  taken  from  one  or  more  of  the  a'bnormal  mainly  pistillate, 
naturally  planted  trees. 


OF  THE 

UNIVERSITY 

OF 


VOL.  2]          Mott.— Teratology  in  Calif ornian  Willows.  197 

the  normal  in  a  manner  very  much  similar  to  that  seen  in  the 
various  other  cases  of  staminody  of  the  pistil  in  Salix  which 
are  described  in  the  literature.  The  stipe  is  usually  lengthened, 
filament-like  (or  like  two  filaments  fused)  ;  the  ovary  is  short- 
ened; the  style  and  stigma  are  more  or  less  nearly  lost;  the 
posterior  suture  is  prematurely  ruptured;  and  two  anthers, 
either  perfectly  developed  or  more  or  less  rudimentary,  are  dis- 
closed, one  on  each  side,  borne  on  the  inner  wall  of  the  ovary; 
a  reduced  number  of  ovules  are  borne  on  the  median,  parietal 
placentae;  pollen  is  produced  and  discharged;  the  exposed 
ovules  soon  shrivel  and  fall.  Probably  a  complete  series  of 
forms  from  pistil  to  stamens  could  be  worked  out  as  in  the  case 
of  the  converse  change. 

The  abnormal  occurrence  of  bisexual  flowers  with  normally 
developed  pistil  and  stamens  (perfect  flowers)  is  a  striking 
peculiarity  of  this  tree  (see  pi.  20,  fig.  20).  The  general  appear- 
ance of  these  flowers  suggests  (if  one  considers  them  with  the 
usually  accepted  theories  of  descent  in  mind),  that  their  occur- 
ence  is  due  to  reversion,  and  hence  that  the  hermaphroditism 
in  them  is  of  a  different  sort  from  the  hermaphroditism  seen  in 
the  monstrous  bisexual  organs  occurring  either  on  this  tree  or 
on  the  one  first  described. 

These  perfect  flowers  are  very  numerous  (though  a  pre- 
ponderance of  normal  pistillate  flowers  on  the  tree  as  a  whole 
shows  it  to  be  predominantly  female),  and  they  are  fairly  uni- 
form in  type,  consisting  usually  of  a  pistil  between  two  stamens, 
the  latter  standing  right  and  left  (transverse).  The  pistil  and 
stamens  are  in  full  bloom  (in  "anthesis")  at  different  times 
(flower  "dichogamous"),  as  is  very  commonly  the  case  in  per- 
fect flowers.  And  here  the  stamens  are  in  anthesis  last ;  that 
is,  the  anthers  do  not  shed  their  pollen  until  the  stigma  is  no 
longer  receptive  (dichogamy  " p rot ogy nous").  The  figure  (20) 
shows  that  the  filaments  of  the  two  stamens  are  still  short,  and 
the  pollen-sacs  unopened,  whereas  the  pistil  is  ready  for  pollina- 
tion. This  difference  in  time  of  anthesis  of  pistil  and  stamens 
in  the  perfect  flowers  is  rendered  more  striking  by  the  fact  that 
there  are  other  stamens  on  this  tree,  often  on  the  same  catkins 


198  University  of  California  Publications.  [BOTANY 

with  perfect  flowers,  yet  usually  occurring  in  purely  male 
flowers,  which  are  in  anthesis,  with  lengthened  filaments,  about 
the  same  time  that  the  pistils  are  in  anthesis.  Sometimes  the 
two  stamens  of  a  perfect  flower,  though  normal  in  form,  are 
small  and  pale  as  though  not  fully  restored. 

Again  there  is  suggestion  of  reversion  to  an  ancestral  form 
of  flower  in  the  fact  that  there  is  a  tendency  on  this  tree, 
especially  in  the  perfect  flowers,  toward  the  establishment  of 
two  extra  glands,  standing  transverse,  in  addition  to  the  usual 
one  or  two  median  (fig.  20  does  not  show  the  extra  glands). 
These  glands  are  referred  to  later  as  possible  vestiges  of  an 
ancestral  perianth.  (They  are  also  described  somewThat  more 
completely  later  on.) 

The  male  flowers  mentioned  as  occurring  on  this  hybrid 
tree  are  rather  common.  There  are  usually  two  or  three  stamens 
in  a  flower,  though  as  high  as  five  occur.  The  supposed  parent 
species  are  tri-  and  pentandrous  respectively. 

Numerous  variations  of  the  types  of  flowers  mentioned 
occur.  Roughly  classifying  all  the  sorts  observed  on  the  tree 
we  have  (1)  normal  pistil  (a)  alone,  (&)  between  two  short* 
stamens;  (2)  monstrous  pistil  (a)  alone,  (b)  between  two  short 
stamens,  (c)  with  one  longf  stamen;  (3)  one  long  stamen  and 
one  short  stamen;  (4)  one  long  stamen  between  two  short 
stamens;  (5)  one  to  five  long  stamens;  (6)  two  long  stamens 
each  with  a  distinctly  double  anther;  (7)  perfectly  developed 
ovary  and  anther  on  the  end  of  one  stalk.  Doubtless  other  com- 
binations and  forms  could  be  found. 

The  aments  are  very  diverse,  as  in  the  case  of  the  Salix  lasio- 
lepis.  The  majority  are  strictly  pistillate,  but  yet  very  many 
are  almost  wholly  of  perfect  flowers,  others  are  purely  staminate, 
and  still  others  are  made  up  chiefly  of  the  flowers  consisting  of 
the  monstrous  organs.  Then  there  are  catkins  that  combine  in 
various  proportions  all  the  types  of  flowers  described. 


*  Short :  that  is,  of  the  set  which  are  late  in  developing  to  shed  their 
pollen. 

t  Long :  that  is,  of  the  set  which  are  early  in  developing  to  shed  their 
pollen. 


VOL.  2]          Mott.— Teratology  in  Calif ornian  Willows.  199 

Branchlets  show  the  same  diverseness,  many  being  largely 
female,  some  chiefly  male,  some  mainly  of  perfect  flowers,  others 
pronouncedly  of  monstrous  organs. 

An  interesting  feature  of  the  teratology  of  this  tree  is  that 
in  this  third  spring  season  the  abnormality  of  the  flowers  is 
more  general  over  the  tree  than  it  was  in  the  first  season.  (The 
tree  was  not  closely  observed  the  second  season).  The  first 
spring,  the  lower  part  of  the  tree  (the  tree  being  about  forty 
feet  tall,  and  of  a  spreading  habit)  was  most  affected,  the  upper 
part  being  but  little  abnormal.  This  spring  the  lower  part  of 
the  tree  is  somewhat  more  affected  than  when  first  noticed,  and 
the  upper  part  has  become  about  as  much  abnormal  as  the  lower. 
Close  to  this  tree  are  two  other  individuals,  apparently  of  the 
same  cross,  and  both  also  mainly  pistillate,  the  three  having  been 
planted  at  the  same  time  as  cuttings;  and  these  others  have 
shown  something  of  the  same  seasonal  increase  of  abnormality 
that  the  first  has  shown.  (To  distinguish  between  the  three 
trees,  the  first  described  may  be  referred  to  as  the  north  tree, 
the  other  two  as  middle  and  south  respectively.)  The  first 
spring  the  middle  and  south  trees  were  not  noticeably  abnormal 
in  any  degree,  though  they  were  carefully  examined.  This 
third  spring,  the  middle  tree  shows  abnormality  in  a  small  pro- 
portion of  the  aments  (the  irregularities  being  of  the  same  gen- 
eral description  as  those  shown  by  the  north  tree)  and  the  south 
tree  has  borne  one  bisexual  flower  with  normal  organs, — a  pistil 
and  one  stamen.  A  thorough  search  over  this  south  tree  failed 
to  reveal  any  other  abnormal  flowers. 

An  irregularity  in  this  one  bisexual  flower  on  the  south  tree, 
which  is  rather  striking  if  one  is  looking  for  evidence  of  rever- 
sion, is  the  presence  of  two  small  lateral  glands,  since  the  normal 
one  or  two  median  glands  in  Salix  are  quite  commonly  con- 
sidered as  vestiges  of  an  ancestral  perianth,  and  here  apparently 
is  indication  of  two  perianth  phyllomes  which  are  normally  en- 
tirely missing.  The  other  flowers  on  the  tree  had  but  a  pos- 
terior gland. 

It  seems  rather  natural  that  restoration,  in  some  degree,  of 
lost  perianth  phyllomes  should  accompany  the  restoration  of  lost 


200  University  of  California  Publications.  [BOTANY 

sexual  phyllomes.  Many  of  the  flowers  also  on  the  north  tree, 
especially  the  bisexual  ones  with  normal  organs,  had,  as  noted 
above,  more  or  less  evident  lateral  glands. 

Some  further  notes  as  to  the  glands  on  these  three  hybrid 
scion  trees  are  of  interest  as  bearing  upon  the  question  of  what 
are  the  species  represented  in  the  trees  (the  descent,  however, 
being  largely  judged  from  other  facts).  On  the  north  tree 
there  are  usually  two  median  glands,  one  anterior  and  one  pos- 
terior, to  each  flower.  (The  glands  are  small  yellow  thallus- 
like,  or  flattened,  bodies  at  the  base  of  the  sexual  organs,  e  and 
/,  pi.  20,  fig.  20).  Now,  of  the  probable  parents,  Salix  lasiandra, 
in  this  locality,  has  commonly  only  the  posterior  gland  in  the 
flowers  of  both  sexes,  and  Salix  babylonica  has  only  the  posterior 
in  the  female  flower,  but  both  anterior  and  posterior  in  the  male. 
The  north  tree,  then,  which  is  quite  largely  bisexual,  seems  to 
follow  the  male  babylonica  in  having  two  median  glands.  It 
should  be  mentioned  here  that  a  male  Salix  lasiandra  found  this 
season,  has  quite  uniformly  four  glands  to  the  flower,  though 
the  tree  is  otherwise  apparently  normal,  and  hence  this  hybrid 
mother  tree  in  showing  a  tendency  to  four  glands,  may  be  merely 
exhibiting  an  occasional  normal  lasiandra  character,  and  not  be 
giving  evidence  of  reversion  in  the  flower  in  this  particular.  (A 
flower  with  the  four  glands  from  this  Salix  lasiandra  is  shown 
in  fig.  22,  pi.  19.  The  five  stamens  and  the  bract  have  been  cut 
off).  The  middle  hybrid  tree,  which  has  a  majority  of  its 
flowers  pistillate,  has  commonly  only  the  posterior  gland,  like 
Salix  lasiandra  and  the  female  babylonica;  and  the  south  tree, 
which  has  all  its  flowers  pistillate,  excepting  the  one  bisexual 
flower  with  the  two  small  lateral  glands  has  aside  from  this  one 
flower  only  the  posterior  gland  throughout. 

TENDENCY  TO  ABNORMALITY  PRESENT  IN  THE  SCIONS  HYBRID, 
SEEMINGLY  BY  PERPETUATION  FROM  PARENT  STOCK;  PROBABLE 
SOURCE  OF  THE  SCIONS,  AND  EVIDENCE  AS  TO  THE  TWO  ORIGINAL 
PARENT  SPECIES. — The  abnormalities  in  these  three  scion-propa- 
gated trees  being  apparently  not  due  to  seasonal  influences  nor 
to  local  conditions  (which  are  normal),  nor  to  such  agencies  as 


VOL.  2]          Mott.— Teratology  in  Calif ornian  Willows.  201 

insects  or  injuries  (for  the  trees  appear  perfectly  healthy  and 
sound),  some  effort  was  made  this  spring  to  find  whether  they 
came  from  abnormal  stock,  and,  as  stated  above  in  a  foot-note, 
their  immediate  origin  was  traced  with  considerable  certainty 
to  some  abnormal  mainly  pistillate  trees  in  a  naturally  grown 
mixed  grove  of  willows  near  by.*  With  the  latter  are  some 
mainly  staminate  abnormal  individuals  of  the  same  cross. 
These  naturally  grown  trees  show  the  same  general  characters, 
both  normal  and  abnormal,  that  mark  the  scion-propagated 
trees.  But  there  are  some  abnormal  features  (some  of  which 
are  mentioned  later),  which  are  rather  more  common  here  than 
on  the  scions,  and  there  is  some  noticeable  variation  from  the 
scions  in  the  relative  prominence  of  the  different  abnormal 
features.f  For  example,  the  tendency  to  four  glands  is  con- 
siderably more  evident  here.  Various  combinations  of  the  four 
glands  occur  in  a  flower,  and  they  may  be  quite  distinct,  or  more 
or  less  united  by  their  bases  into  a  ring  encircling  the  sex-organs, 
like  a  miniature  sympetalous  corolla. 

The  hybrid  origin  of  all  these  abnormal  trees — such  descent 
seeming  quite  certain — is  of  interest  as  being  very  likely  the 
cause  of  the  teratology  which  marks  them,  hybrids  in  general 
being  prone  to  show  abnormalities,  especially  in  the  flowers. 
The  scion  trees  were  first  taken  for  lasiandras,  since  they  re- 
semble that  species  more  than  any  of  the  other  native  species, 
but  later  some  planted  exotic  pistillate  babylonicas,  or  weeping 
willows,  were  found  near  one  edge  of  the  natural  grove  of  wil- 
lows, and  these  are  probably  the  original  maternal  stock  of  all 
the  abnormal  trees,  while  native  lasiandras,  which  are  common 
in  the  neighborhood,  are  probably  the  paternal  stock.  The 
babylonicas  were  planted  some  fifty  years  ago  to  mark  some  old 
Spanish  graves,  and  their  flowers  were  probably  pollinated  at 
some  time  from  lasiandras  and  then  seeds  from  them  blew  over 
and  gave  rise  to  the  abnormal  trees  in  the  grove.  The  grove 

*  Normal  trees  in  the  grove  are :  many  Salix  lasiolepis  Benth.,  a  few 
S.  lasiandra  Benth.,  and  a  few  S.  laevigata  Bebb. 

t  The  matter  of  whether  scions  from  these  hybrid  willows  will  per- 
petuate the  floral  abnormalities,  in  some  measure,  as  seems  almost  certainly 
to  have  occurred,  is  being  tested  in  the  University  gardens  with  some  cut- 
ting from  one  of  the  trees.  There  are  no  flowers  this  first  season. 


202  University  of  California  Publications.  [BOTANY 

came  into  notice  only  about  twenty-five  years  ago,  and  the  trees 
forming  it  were  still  quite  small  when  the  three  cuttings  were 
taken. 

The  probable  origin  of  these  abnormal  trees  by  the  crossing 
of  these  particular  species  is  indicated  in  the  trees  themselves 
by  their  showing  many  characters  intermediate  between  the 
normal  characters  of  8.  lasiandra  and  8.  babylonica, — as  to 
habit,  number  of  stamens,  number  of  glands  (as  noted  above), 
and  so  on.  It  is  true  that  the  common  presence  in  the  perfect 
flowers  of  two  stamens  might  suggest  that  the  male  parent  was 
of  some  such  native  diandrous  species  as  Salix  lasiolepis  Benth ; 
for,  of  the  two  assumed  parent  species,  Salix  lasiandra  has  5-4 
stamens  (for  this  region),*  and  Salix  babylonica  three,  and, 
furthermore  the  purely  male  flowers  on  the  abnormal  trees  are 
most  frequently  of  two  or  three  stamens;  but  yet  the  abnormal 
trees  have  not  uncommonly  as  high  as  five  stamens  in  the  purely 
male  flowers,  like  normal  8.  lasiandra,  and  besides  they  have 
many  characters,  aside  from  the  number  of  stamens,  which  seem 
to  preclude  the  supposition  of  parentage  on  one  side  by  any  of 
the  native  species  other  than  8.  lasiandra.j 

It  may  be  noted  in  connection  with  this  matter  of  parentage, 
that  Salix  babylonica  has  been  frequently  reported  as  abnormal 
in  the  flowers. 

ON  SOME  NATURALLY  GROWN  INDIVIDUALS  APPARENTLY  RE- 
LATED TO  THE  SCION  TREES: SEPARATION,  PARTIAL  TO  COMPLETE, 

OF   THE   TWO    CARPELS   OF   THE  PISTIL  WITHOUT   BISEXUALISM,   AND 

ALSO  MULTIPLICATION  OF  CARPELS. — These  two  irregularities  are 
the  most  striking  among  those  referred  to  above  as  occurring 
commonly  on  some  of  the  abnormal  trees  in  the  grove  and  not 
to  any  extent  on  the  scion  trees.  The  abnormalities  are  most  in 
evidence  on  the  mainly  pistillate  trees.  Cases  illustrative  of 

*  Jepson,  Flora  of  Western  Middle  California. 

t  A  case  of  the  occurrence  of  willows  which  were  considered  to  be 
Salix  lasiandra  X  babylonica,  from  a  cross  between  native  staminate 
lasiandras  and  planted  exotic  pistillate  babylonicas  was  reported  by  Ander- 
son (1890)  from  Santa  Cruz,  Calif.  These  trees  were  described  on  account 
of  being  monoecious,  staminate  and  pistillate  flowers  occurring  together  on 
the  catkins,  but  none  of  the  other  abnormalities  that  are  present  on  these 
Berkeley  hybrids,  are  mentioned. 


VOL.  2]          Mott.— Teratology  in  Calif  or  nian  Willows.  203 

these  two  general  manifestations  of  abnormality  have  been  fre- 
quently reported  by  writers,*  so  mere  mention  of  some  of  the 
forms  in  point  found  on  our  trees  will  be  made.  There  are 
found  then  here:  (1)  pistils  showing  various  degrees  of  separa- 
tion of  the  carpels;  (2)  two  distinct  unicarpellary  pistils  re- 
sembling miniature  pea-pods  in  one  flower;  (3)  three  such  pistils 
in  one  flower;  (4)  a  normal  bicarpellary  and  a  unicarpellary 
pistil  in  one  flower;  (5)  two  perfectly  developed  and  distinct 
bicarpellary  pistils  in  one  flower.  Sometimes  the  unicarpellary 
pistils  were  monstrous  in  being  bisexual,  like  the  bicarpellary 
pistils  described  above,  a  more  or  less  malformed  anther  bulging 
out  at  the  suture  between  the  carpel  margins. 

Bearing  of  the  Abnormalities  Described  on  Some  Moot 
Points  Regarding  the  Salix  Flower. 

The  notes  following  here  are  based  mainly  on  the  usually 
accepted  general  theories  of  evolution.  Most  comment  of 
writers,  suggested  by  similar  abnormalities  in  Salix,  has  been  so 
made,  though  the  conclusions  to  which  different  writers  have 
inclined  have  differed.  A  more  recent  fundamental  theory, 
under  which  such  abnormalities  may  be  interpreted,  is  referred 
to  later. 

WAS  THE  ANCESTRAL  FLOWER  BISEXUAL  OR  UNISEXUAL?       This 

question  is,  perhaps,  the  one  most  commonly  discussed,  and  the 
answer  to  it  seems  to  depend  mainly  upon  the  interpretation 
one  makes  of  the  present  sporadic  instances  of  hermaphroditism 
in  the  flower.  In  fact,  it  is  generally  the  encountering  of  cases 
of  hermaphroditism  that  raises  this  question  about  the  ancestral 
flower. 

On  our  trees  there  seem  to  be  two  distinct  sorts  of  hermaphro- 
ditism. On  all  the  trees  there  are  (1)  the  flowers  with  the 
monstrous  bisexual  phy Homes,  and  (2),  especially  on  the  hybrid 
tree  first  described,  the  flowers  with  the  normally  developed  pistil 
and  stamen.  On  this  latter  tree,  the  two  kinds  of  androgyny 


*  Von   Seemen    (1886  and   1895),   in   particular,   has  pictured   and   de- 
scribed many  abnormal  forms  coming  under  these  two  heads. 


204  University  of  California  Publications,  [BOTANY 

are  so  associated  as  apparently  to  show  clearly  that  they  are 
distinct  phenomena ;  and  the  perfect  flowers  with  normal  organs, 
as  noted  in  describing  them,  certainly  show  many  of  the  gen- 
erally accepted  signs  of  reversion,  entirely  aside  from  any  con- 
sideration of  the  monstrous  bisexual  organs,  and  hence  they 
seem  to  indicate  that  the  ancestral  flower  was  bisexual. 

Some  writers  (Camus,  1889,  and  Von  Seemen,  1886  and 
1895)  have  inclined  to  the  belief  that  the  perfect  flowers  often 
seen  have  come  about  by  sexual  change  of  part  of  the  phyl- 
lomes  of  an  always  unisexual  flower,  and  point  to  such  cases 
of  complete  substitution  of  one  sexual  organ  for  the  other  as 
seen  in  our  Salix  lasiolepis  series,  to  show  the  possibility  of  the 
origin  of  bisexual  flowers  with  normal  organs  in  such  a  way;* 
and  on  any  polyandrous  male  individual  such  origin  does  seem 
possible,  for  there  are  sufficient  phyllomes  there  in  one  flower 
for  both  stamens  and  pistil.  But  on  a  pistillate  individual, 
with  its  two  phyllomes  to  a  flower,  such  as  our  hybrid,  we 
should  have  to  conceive  of  first  a  multiplication  of  phyllomes  and 
then  a  development  of  them  as  stamens  instead  of  as  carpels. 
Multiplication  of  phyllomes  is  a  common  phenomenon,  it  is 
true; — the  polyandry  in  some  species  of  willowsf  and  the  more 
extreme  cases  of  polyandry  in  the  other  genus  of  the  family, 
the  poplars,|  doubtless  arose  by  reduplication,  and  the  extra 
unicarpellary  and  bicarpellary  pistils  on  some  of  our  mainly 
pistillate  hybrids  are  probably  to  be  accounted  for  in  this  way ; 
but  it  seems  far-fetched,  in  the  case  of  the  perfect  flowers  on 
our  predominantly  pistillate  hybrid,  which  flowers  consist  rather 
uniformly  of  pistil  between  two  stamens,  to  assume  that  these 
two  lateral  stamens  are  present  through  reduplication  of  phyl- 

*  Von  Seemen  and  others,  interpreting  in  this  way  the  apparent  mutual 
convertibility  of  pistil  and  stamens,  have  held  that  pistils  and  stamens  in 
Salix  are  morphologically  identical.  But  if,  however,  there  are  sufficient 
facts  which  can  be  interpreted  as  indicating  that  the  ancestral  flower  was 
bisexual,  there  would  seem  to  be  no  more  reason  for  considering  "the  male 
and  female  sex-organs  morphologically  equivalent"  here  than  in  flowers  in 
general. 

f  As  high  as  twenty  stamens  occurring  in  some  species,  according  to  Pax 
(in  Engler  and  Prantl's  Pflanzenfamilien). 

\  As  many  as  thirty  stamens  being  sometimes  present  here,  according  to 
Pax;  and  even  seventy-one  according  to  Jepson,  Flora  of  Western  Middle 
California,  2nd  ed.,  MS. 


VOL.  2]          Mott— Teratology  in  Calif ornian  Willows.  205 

lomes,  and  then  that  they  are  stamens  by  conversion.  Although 
monstrous  bisexual  organs  occur  on  this  tree,  even  in  the  same 
catkin  with  the  perfect  flowers,  these  two  lateral  stamens  are 
not  found  connected  by  any  intermediate  forms,  in  the  same 
position  in  the  flower,  with  carpels.  They  look  like  restored 
organs, — original  microsporophylls.  The  bisexual  bodies  that  do 
occur,  on  the  contrary,  seem  always  to  be  more  or  less  changed 
fo'carpellary  pistils,  standing  in  the  normal  position  of  the  pistil. 
They  may  even  occur  in  the  place  of  the  pistil  in  otherwise  per- 
fect flowers. 

Further  facts  indicating  reversion  here,  and  hence  indicat- 
ing once  perfect  flowers,  are,  that  the  perfect  flowers  are  pro- 
togynous,  that  their  stamens  are  sometimes  small  and  pale,  as 
though  incompletely  restored,  that  the  perfect  flowers  are  numer- 
ous and  of  a  fairly  uniform  type,  that  there  is  a  tendency  to  a 
concomitant  restoration  of  glands,  which  seem  to  be,  with  the 
normal  glands,  perianth  vestiges,  and  that  there  are  only  two 
stamens  in  these  perfect  flowers,  as  though  by  restoration  of  an 
original  number  (as  opposed  to  a  later  tendency  to  multipli- 
cation of  stamens  in  the  family). 

Miiller  believed  the  ancestor  of  the^present  unisexual  flower 
was  bisexual,  and  was  led  to  this  belief  by  consideration  of  the 
same  evidence  of  possible  change  of  organs  (as  seen  in  our  Salix 
lasiolepis  series)  which  has  led  others  to  believe  the  present  occa- 
sional bisexual  flower  with  normal  organs  is  derived  from  an 
ancestral  unisexual  flower.  Since  stamens  can  thus  change  com- 
pletely to  pistils,  and  vice  versa,  he  concluded  that  the  modern 
unisexual  flower  developed  one  way  or  the  other  from  a  bisexual 
flower.  But  if  that  be  true,  why  have  the  normal  pistillate  flowers 
and  the  diandrous  male  flowers  only  two  phyllomes  each?  Evi- 
dently we  must  assume  suppression  there  of  the  phyllomes  of 
the  lost  sex  (suppression  being  in  fact  a  common  occurrence, 
as  noted  later),  and  hence  why  not  assume  that  there  has  oc- 
curred uniformly  a  suppression  of  the  phyllomes  of  one  sex  or 
the  other  in  an  ancestrally  bisexual  flower,  to  produce  the  mod- 
ern flower,  and  then  that  the  increased  number  of  phyllomes 
in  polyandrous  species  has  occurred  uniformly  by  reduplication  ? 


206  University  of  California  Publications.  [BOTANY 

This  monstrosity  of  bisexualism  occurring  in  one  organ  seems 
capable  of  a  double  interpretation,  and  apparently  is  one  of 
those  numerous  teratological  phenomena  that  must  be  consid- 
ered as  of  no  morphological  significance.  As  noted  above,  this 
hermaphroditism  shown  in  the  monstrous  organs  seems  to  be  of 
a  sort  distinct  from  that  exhibited  in  the  perfect  flowers  with 
normal  organs. 

Aside  from  the  evidence  that  has  been  noted,  which  it  seems 
may  be  accepted  as  showing  that  the  ancestral  flower  was  bisex- 
ual although  it  is  teratological  evidence,  there  is  the  additional 
strong  support  of  the  theory  of  original  bisexualism  in  the 
great  similarity — in  most  species  practical  identity  aside  from 
the  sexual  organs — of  the  male  and  the  female  aments,  in  all 
characters, — shape,  size,  coloring,  position  on  the  branches,  char- 
acter of  the  individual  bracts,  degree  of  hairiness,  etc.  In  the 
other  amentiferous  families  with  unisexual  flowers,  such  as  the 
oaks,  walnuts,  hazels,  and  even  birches  and  alders,  there  is 
a  very  great  difference  between  the  floral  clusters  of  the  two 
sexes.  In  such  cases  it  might  be  difficult  to  imagine  how  a  uni- 
sexual flower  could  be  the  result  of  suppression  of  one  of  the 
sexes  in  a  perfect  flower.  But  in  Salix,  unisexualism  by  such  a 
process  is  easy  to  conceive. 

That  unisexual  flowers  may  arise  by  abortion  of  one  sex,  an- 
droecium  or  gynaeceum,  is  shown,  it  is  commonly  considered, 
beyond  doubt,  in  such  a  case  as  that  of  the  polygamous  genus, 
Acer,  the  maples.  Here  there  are  sometimes  bisexual  flowers 
to  be  found  with  organs  of  both  sexes  normal ;  in  other  flowers, 
the  organs  of  one  sex  have  become  vestigial,  though  they  are 
still  recognizable  as  being  remnants  of  such  organs,  and  in  still 
others,  the  organs  of  one  sex  have  suffered  complete  obliteration. 

That  suppression  is  common  in  the  willow  family,  Pax  * 
thinks  is  shown  in  the  reduction  of  the  broad  disk  of  Populus 
to  the  usual  one  or  two  small  glands  in  Salix.  Evidently  he  con- 
siders both  disk  and  gland  as  reduced  perianth. 

On  the  whole,  there  seem  good  reasons  for  believing  that  the 
ancestral  Salix  flower  was  bisexual. 


*  In  Engler  and  Prantl's  Pflanzenfamilien. 


VOL.  2]          Mott.— Teratology  in  Calif ornian  Willows,  207 

ARE  THE  FLORAL  GLANDS  VESTIGES  OF  AN  ANCESTRAL  PERIANTH  ? 

— The  glands  of  Salix  have  been  held  by  several  writers  to  be 
vestigial  perianth  of  a  once  bisexual  flower.  Velanovsky  (1904), 
especially,  has  recently  worked  out  a  ground  plan  for  a  com- 
plete flower,  basing  his  contentions  as  to  perianth  on  some  tera- 
tological  material  of  Salix  aurita  L.  The  flowers  were  bisexual, 
and  the  posterior  gland  throughout,  according  to  his  interpre- 
tation, was  divided  into  two  somewhat  leaf-like  bodies,  which, 
further,  had  been  moved  around  so  as  to  stand  transverse.  He 
then  restores  a  lost  posterior  gland  to  match  the  anterior  one, 
and  has  an  ideal  dimerous  perianth,  as  in  Myrica  and  Juglans, 
which  some  consider  as  standing  next  in  relationship  to  Salix. 

The  occurrence  of  four  glands  together  as  noted  above,  a 
fact  which  apparently  has  not  been  recorded  before  in  Salix. 
would  seem  to  make  possible  a  much  simpler  restoration  of  the 
same  perianth.  The  four  glands  might  be  vestiges  of  four  peri- 
anth phyllomes,  in  the  original  positions  of  those  phyllomes. 
Then  to  get  the  modern  normal  flower,  with  its  posterior  gland 
and  sometimes  also  anterior  gland,  we  have  only  to  conceive 
that  the  other  glands,  or  phyllomes,  have  been  entirely  sup- 
pressed. In  the  reversion  to  hermaphrodite  flower  in  our  hybrid, 
there  seems  to  be  a  tendency  to  restore  these  lost  glands. 

SUPPOSING  THE  ANCESTRAL  FLOWER  WERE  BISEXUAL,  WAS  THE 
NUMBER  OF  STAMENS  TWO  OR  FOUR?  ARE  THE  TWO  CARPELS  OF 

THE  PISTIL  NOW  MEDIAN  OR  TRANSVERSE. — Velanovsky  arguing 
for  a  once  perfect  flower,  concludes  that  it  had  four  stamens.  For 
since  the  two  carpels  of  the  pistil  stand  transverse  in  Salix.  and 
since  the  pair  of  stamens  also  in  the  perfect  flowers  of  his  mate- 
rial stand  transverse  (as  in  ours),  to  obtain  a  dimerous  flower 
with  regular  alternation  of  parts,  he  restores  a  median  pair  of 
stamens  and  thus  has  four.  However,  since  the  actually  occur- 
ring perfect  flowers  on  our  hybrids  give  many  indications  of 
reversion  to  the  original  type,  and  these  flowers  have  two  stamens 
each,  perhaps  we  could  as  well  assume,  to  satisfy  alternation, 
that  the  two  stamens  were  once  median  and  were  crowded  around, 
to  stand  transverse,  by  the  closely  appressed  subtending  bracts 


208  University  of  California  Publications.  [BOTANY 

of  the  ament.  Such  displacement  in  general  is  not  uncommon. 
Yet,  again,  is  it  necessary  that  alternation  be  satisfied?  It  is 
frequently  lacking  in  these  ancient  types.  Regularity  in  number 
and  in  alternation  of  parts  is  a  development  of  the  modern 
flower.  It  would  seem  that  we  can  as  well  hold  that  the  original 
number  of  stamens  was  two. 

Pax,*  arguing  for  a  once  perfect  flower,  gives  a  ground  plan 
based  on  the  case  of  abnormality  reported  by  Heinricher  (1883), 
and  shows  in  it  only  two  stamens.  However,  he  represents  the 
carpels  as  standing  median,  and  thus  there  is  alternation  with 
only  the  two  stamens. 

Yet — to  consider  now  whether  the  carpels  are  median  or 
transverse — this  median  position  given  by  Heinricher  seems 
wrong.  He  had  but  a  single  flower,  and  does  not  support  his 
statement  as  to  the  position  of  the  carpels  by  any  evidence.  The 
stipe  of  his  pistil,  too,  is  shown  to  be  abnormally  long,  and  in 
such  cases  it  is  difficult  to  make  sure  of  the  position  of  the  car- 
pels. The  formation  of  a  perfectly  normal  pistil  from  two  trans- 
verse stamens,  as  seen  in  our  8.  lasiolepis  and,  conversely,  the 
median  rupturing  of  a  pistil  to  form  two  transverse  stamens, 
seem  safely  to  be  taken  as  evidence  (as  noted  by  Wichura  as 
long  ago  as  1887),  that  the  carpels  of  the  pistil  stand  transverse. 
This  is  the  position  accepted  for  the  normal  female  flower  by 
Pax,  by  Sargent,  and  by  others,  in  descriptions  of  the  genus 
(though  Brendel  [1880]  appears  to  think  the  median  position 
generally  held). 

Hegelmaierf  is  led  to  believe  in  the  transverse  position  of 
the  carpels  by  a  study  of  the  development  of  the  pistil  in  the 
bud.  On  the  other  hand,  an  examination  of  the  winter  flower 
buds  of  Salix  lasiolepis  seems  not  to  bear  out  Hegelmaier's  view, 
— however  much  other  evidence  does.  The  pistil  appears  as  a 
small  medianly  compressed,  urn-shaped  protuberance,  bilabiate 
at  the  apex,  with  the  lips  anterior  and  posterior  seeming  to  cor- 
respond to  carpel  apices.  There  are  even  two  slight  lateral 

*  In  Engler  and  Prantl's  Pfianzenfamilien,  III,  1,  p.  31. 

f  tiber  Bliitenentwickelungen  bei  den  Salicineen.  Jahreshef  te  des  Ve- 
reins  fur  Vaterlandischer  Naturkunde  in  Wiirttemberg.  Stuttgart,  1880. 
XXXVI,  pp.  204-244,  Tafn.  Ill,  u.  IV. 


VOL.  2]          Mott— Teratology  in  Calif  ornian  Willows.  209 

grooves,  appearing  to  indicate  lateral  sutures,  and  here  are  only 
very  slight  indentations  in  the  middle  of  each  lip.  This  sug- 
gests strongly  that  the  carpels  are  anterior  and  posterior  (me- 
dian), and  that  the  valves  of  the  capsule,  which  are  median, 
correspond,  therefore,  to  the  carpels.  However,  such  a  series 
of  forms  connecting  a  pair  of  transverse  stamens  and  a  normal 
pistil,  as  shown  in  the  drawings  here  of  Salix  lasiolepis,  and 
also  the  converse  phenomenon,  as  indicated  in  the  figure  from 
the  hybrid,  would  seem,  even  though  the  evidence  is  terato- 
logical,  to  make  the  contention  for  the  transverse  position  of  the 
carpels  prevail. 

This  view  is  strengthened  by  the  median  position  of  the  two 
placentae,  since  ovules  are  commonly,  though  by  no  means 
always,  marginal  on  the  carpels.  On  the  other  hand,  the  trans- 
verse dehiscence  of  the  capsule  in  Salix  might,  by  itself,  lead 
one  to  conclude  that  the  sutures  are  transverse — and  hence  the 
carpels  median — for  dehiscence  of  unilocular  capsules  is  per- 
haps more  commonly  at  the  sutures  than  at  the  midribs;  but 
in  some  such  capsules,  as  in  the  tricarpellary  fruit  of  Viola,* 
the  splitting  does  occur  at  the  midribs,  and  in  septate  capsules, 
dehiscence  is  quite  commonly  loculicidal. 

The  balance  of  evidence  seems  to  show  that  the  carpels  in 
Salix  stand  transverse ;  and,  yet,  at  the  same  time,  the  necessity 
for  assuming  more  than  two  stamens  in  the  ancestral  flower  seems 
questionable. 

Some  Possible  Causes  of  the  Abnormalities  Described. 

Any  attempt  to  discover  the  ultimate  cause  of  such  abnorm- 
alities as  found  in  the  two  willows  here  described  would  doubt- 
less entail  an  explanation  of  all  the  manifold  teratological  phe- 
nomena that  occur  throughout  the  plant  kingdom,  and  involve 
one,  as  noted  at  the  outset,  in  still  wider  biological  problems. 
The  proximate  cause  in  the  case  of  the  Salix  lasiandra  X  baby- 
lonica,  at  least  (if  we  confine  ourselves  to  consideration  of  the 
fairly  well  established  causes  of  teratology),  seems  to  be  the 

*  Sachs,  Text  Book  of  Botany. 


210  University  of  California  Publications.  [BOTANY 

crossing  of  species.  Considerable  evidence  points  to  this. — Wil- 
lows in  general  are  very  prone  to  this  intermingling  ("Wichura 
by  repeated  crossing  obtaining  a  hybrid  combining  no  less  than 
six  species).  A  large  proportion  of  the  abnormal  cases  reported 
for  Salix  are  of  hybrids.  In  general,  "  malformations  and  odd 
forms  are  apt  to  appear  in  hybrids,  especially  in  the  flowers;"* 
"monstrous  or  abnormal  forms  of  floral  organs  are  much  more 
common  in  hybrids  than  in  individuals  of  pure  descent, '  'f  Mil- 
lardet,±  in  extended  experiments  with  strawberries,  found  that 
many  hybrid  descendants  bore  perfect  flowers  instead  of  uni- 
sexual with  the  male  and  female  on  different  plants. 

It  seems  quite  possible  that  many  of  the  cases  of  abnormal 
willows  reported  under  a  simple  specific  name  would  prove,  on 
careful  investigation,  to  be  hybrids,  as  happened  with  our  8. 
lasiandra  X  babylonica.  This  might  be  true  of  our  8.  lasiolepis, 
though  it  appeared  pure.§  In  general,  it  is  true,  hybrids  are 
more  or  less  intermediate  in  characters  between  the  parents,  yet 
it  is  not  uncommon 1 1  for  the  offspring  to  resemble  one  parent 
exclusively.  Or,  in  all  outward  characters,  as  of  habit,  foliage, 
and  fruit,  offspring  may  resemble  one  parent,  while  in  hardi- 
ness, vigor,  resistance  to  disease,  fertility,  etc.,  they  may  follow 
the  other.  In  such  cases  there  would  still  be  a  mingling  of  blood, 
as  it  were,  to  cause  disturbance  internally  in  the  adjustment  of 
vital  forces  that  had  been  reached  in  each  species,  and  such  dis- 
turbance might  easily  lead  to  manifestation  in  morphological 
abnormalities. 

Sometimes  teratological  phenomena  seem  to  be  due  to  change 
in  nutrition,  rich  or  unusually  abundant  food,  cultivation,  or 
other  change  of  normal  conditions  which  occasions  such  a  physio- 

*  Bailey,  Plant  Breeding,  p.  231. 

t  Focke,  quoted  by  Swingle  and  Webber,  Hybrids  and  their  Utilization 
in  Plant  Breeding.  Yearbook  of  the  U.  S.  Dept.  of  Agric.  for  1897,  p.  410. 

;  L.  c.,  p.  398. 

§  The  native  willows  of  the  region,  aside  from  Salix  lasiolepis,  which 
might  possibly  have  given  rise  to  this  tree  by  crossing  are  S.  lasiandra 
Benth.,  S.  laevigata  Bebb,  and  S.  Nuttalli  Sarg.  var.  brachystachys.  Sarg. 
A  few  introduced  species,  like  S.  babylonica,  have  been  planted  to  some 
extent  in  the  locality.  In  general  appearance,  however,  this  abnormal  tree 
was  identical  with  many  individuals  of  S.  lasiolepis  with  which  it  was  grow- 
ing. All  these  trees  were  naturally  planted. 

II  L.  6:,  p.  396. 


VOL.  2]          Mott. — Teratology  in  Calif  ornian  Willows.  211 

logical  disturbance  as  suggested  above.  In  these  two  instances 
of  the  willow,  however,  there  is  no  indication  of  any  such  excit- 
ing cause,  for  trees  growing  near  by,  under  identical  conditions, 
are  perfectly  normal. 

The  author  of  one  of  the  articles  on  Salix, — writing  also  of 
Carex — (Hampe,  1840),  thought  that  his  observations  showed 
that  dry  conditions  tend  to  produce  change  of  male  organs  to 
female,  and  moist  conditions  the  reserve.  But,  as  shown  (by 
Von  Schlectendal)  in  a  discussion  which  follows  the  article,  this 
explanation  does  not  always  hold.  Our  8.  lasiolepis,  in  fact 
though  changing  to  female,  was  on  a  small  creek  and  none  of 
the  other  adjacent  trees,  in  the  same  conditions,  showed  the 
peculiarity. 

Insects  are  doubtless  responsible  for  some  malformations,  as, 
for  example,  the  willow-roses  and  galls.  Their  sting,  or  punctur- 
ing of  a  growing  shoot  to  deposit  an  egg,  introduces  foreign 
chemicals,  either  directly  with  the  egg,  or  in  the  course  of  devel- 
opment of  the  larva,  into  the  tissues  of  the  plant;  and  this 
causes  a  disturbance  of  the  normal  stimuli  of  growth.  Para- 
sitic fungi,  too,  give  rise  to  disorder  by  chemical  means  and 
they  have  been  known  to  cause  the  production  of  bisexual  flowers 
where  unisexual  are  normal.* 

In  the  case  of  our  S.'lasiandra  X  babylonica,  however,  there 
was  no  notable  evidence  of  insect  action ;  and  in  the  case  of  the 
8.  lasiolepis,  though  there  was  considerable  breeding  of  insects 
on  the  tree,  still  adjacent  trees  were  quite  as  much  affected,  and 
they  were  normal. 

Hybridizing  here  seems  to  be  the  most  likely  explanation  of 
both  abnormalities,  the  monstrous  organs  and  the  perfect  flowers. 

There  may  be  some  elucidation,  if  not  explanation,  of  these 
and  other  teratological  phenomena,  in  Sachs '  ' '  Stoff  und  Form ' ' 
theory,  or,  perhaps  better,  hypothesis,  first  advanced  in  1875, 
and  further  developed  and  explained  in  1892.  f  According  to 
his  idea,  the  plant  produces  a  special  substance  for  each  organ's 
development  out  of  the  common  organic  compounds  which  go 

*  Strasburger :  Biologisches  Centralblatt,  20:  657.     1900. 
t  Sachs '  Gesammelte  Abhandhmgen  iiber  Pflanzen-Physiologie.     II,  pp. 
1159-1231. 


212  University  of  California  Publications.  [BOTANY 

to  form  the  mass  of  all  the  organs.     For  each  organ  there  is  a 
different  substance.     The  amount  of  this  special  substance  may 
be  very  minute  in  proportion  to  the  total  amount  of  tissue  going 
to   form  the   organ.     Under  normal  conditions   each   "special- 
baustoff"  is  conducted  to  a  particular  part  of  a  growing  point 
to  produce  the  organ  normal  there, — stem,  root,  leaf,  stamen, 
pistil,  etc.    If  any  unusual  stimuli  are  introduced  (as  by  cross- 
ing, insects,  change  in  nutrition),  the  attunement  of  forces  in 
the  plant  is  disturbed,  and  there  may  result  a  misdirection  of 
these  special  substances — a  "misleitung  der  Specialbaustoffe  "- 
and  pistil-substance,  for  example,  will  go  to  developing  tissue 
which  would  normally  have  become  stamens,  and  will  cause  the 
production  of  a  pistil  there;  or  stamen-substance  will  be  sent 
astray,  and  will  invade  the  embryonic  tissue  which  could  nor- 
mally have  developed  into  pistil,  and  cause  the  production  of 
stamens  there,  and  so  on.     Or  there  may  be  a  combination,  in 
various  proportions,  of  two  of  these  special  substances,  and  the 
result  will  be,  for  example,  a  dual  organ  partaking  of  the  char- 
acters of  both  pistil  and  stamen,  the  preponderance  of  the  one 
set  of  characters  or  of  the  other  depending  upon  the  proportion 
of  the  special  substances  entering  in. 

Flowers  are  most  likely  to  be  the  region  of  such  disturbances 
owing  to  the  close  association  there  of  several  organs  of  very 
different  nature.  A  very  minute  deviation  of  a  special  sub- 
stance on  its  way  to  produce  one  of  these  organs  is  sufficient  to 
throw  it  into  a  position  which  is  normally  the  sphere  of  action 
of  one  of  the  other  special  substances. 

If  there  is  overproduction,  for  any  reason,  of  any  special 
substance,  then,  under  the  theory,  there  results  a  reduplication, 
or  multiplication,  of  the  appropriate  organ.  And  thus,  for  ex- 
ample, there  come  to  be,  in  many  species  of  Salix,  a  greater 
number  of  stamens  than  the  original  number, — which  we  have 
thought  to  be  two.  The  theory  could  further  be  used,  it  would 
seem,  to  account  for  the  present  occasional  occurrence  of  perfect 
flowers  in  Salix.  Supposing,  for  the  reason  outlined  above,  the 
flowers  were  once  perfect,  the  plant  then  possessed  the  power, — 
and  it  was  active — to  produce  both  the  special  stamen-substance 


ySv 

\ 

I   UNIVERSITY  | 


VOL.  2]          Mott. — Teratology  in  Calif ornian  Willows.  213 

and  the  special  pistil-substance.  Then,  owing  to  the  rise  of  cer- 
tain conditions,  the  power  to  produce  either  the  one  or  the  other 
of  these  special  substances  became  dormant,  and  the  flowers  be- 
came unisexual,  with  male  and  female  flowers  on  separate  trees. 
Finally,  later  conditions,  as  noted  above,  such  as  crossing,  re- 
awaken in  certain  individuals  this  dormant  power,  and  there 
occurs  reversion  to  the  bisexual  form  of  flower. 

The  preceding  discussion  has  been,  of  course,  on  the  basis  of 
the  long  widely-accepted  theories  of  descent  and  of  reversion.  It 
is  well,  perhaps,  finally,  to  recognize  at  least  the  more  recent 
theory  of  the  origin  of  characters  which  is  coming  to  be  favor- 
ably considered  by  many.  This  theory  says,  that  if  the  willow, 
which  certainly  has  not  been  a  complex  organism  since  all  time, 
could  once  develop,  for  example,  the  power  to  produce  perianth 
and  perfect  flowers,  it  could  lose  that  power  completely,  not 
merely  suffer  its  becoming  dormant,  and  then  develop  it  anew, 
as  at  first,  and  the  phenomenon  could  not  be  called  reversion: 
or  it  could  be,  that  our  willow  has  now  developed  these  charac- 
ters for  the  first  time,  and  nothing  is  to  be  concluded  as  to 
whether  or  not  it  ever  possessed  them  before.  And  similarly 
other  abnormalities  herein  noted  would  be  explained. 

Summary.     ,. 

On  a  tree  mainly  staminate,  of  the  diandrous  Salix  lasiolepis 
Benth.,  abnormal  flowers  were  produced  which  show  a  complete 
series  of  forms  in  change  of  a  normal  pair  of  stamens  into  a 
normal  bicarpellary  pistil  (pis.  19,  20,  figs.  2-13). 

X  given  phyllome  is  constant  in  character.  A  pair  usually 
show -equal  change  (e.g.,  pi.  19,  figs.  5  and  9),  but  may  differ 
somewhat  (pi.  19,  figs.  4  and  7).  Through  the  series  the  form  of 
the  phyllome  varies  from  stamen  to  carpel  in  proportion  (with 
some  latitude)  as  its  pollen-sacs  degenerate  and  ovules  supersede 
them.  Pollen-sacs  and  ovules  are  produced  from  the  same  part 
of  the  phyllome  (pi.  19,  figs.  4  and  8)  ;  connective  and  placental 
regions  seem  homologous.  While  phyllomes  are  immature,  an- 
thers, when  present  are  introrse  (pi.  19,  fig.  3)  (though  normally 
in  this  species  extrorse,  pi.  19,  fig.  1  A);  for  the  phyllomes  stand 


214  University  of  California  Publications.  [BOTANY 

with  their  reproductive  faces  together  as  looking  to  the  enclosing 
of  the  ovules.  Dehiscence  of  the  imperfect,  bisexual  ovary  is 
sutural  (pi.  19,  figs.  7  and  8)  ;  phyllomes  remain  intact  (standing 
transverse).  When  pollen-sacs  have  disappeared,  dehiscence  (as 
normally  in  Salix)  is  at  the  midribs, — loculicidal  (valves  median) 
(pi.  20,  fig.  12).  In  the  former  case  dehiscence  is  premature, 
owing  to  swelling  of  the  pollen-sacs  and  their  twisting  to  face 
(as  normally)  the  honey-gland;  in  the  latter  case,  it  is  at  the 
normal  time.  Exposed  ovules  do  not  become  mature  seeds. 

A  rudimentary  ovule  often  causes  an  ovule-appearing  lobe  to 
form  in  the  wall  of  a  pollen-sac  developing  around  and  over  it 
(d  and  e,  pi.  19,  fig.  5)  ;  or,  developing  normally,  the  ovule  may 
stand  protruded  through  the  pollen-sac  (a,  pi.  19,  fig.  5).  Ves- 
tigial pollen-sacs  often  stand  independent  of  each  other,  showing 
the  individuality  of  the  pollen-sac  as  a  microsporangium  (d,  pi. 
19,  fig.  7).  Pollen-sac  vestiges  and  lobes  are  distinguished  from 
ovules  by  their  normal  structural  peculiarities,  and  often  by 
pollen-grains  (pi.  20,  figs.  17-19),  and  ovules  are,  in  the  same 
way,  identified  by  structural  peculiarities  (pi.  20,  fig.  16). 

Monstrous  bisexual  organs  tend  to  make  a  catkin  fall  early ; 
a  very  few  normal  pistils  keep  a  catkin  otherwise  male  or  of 
monstrous  organs  from  falling  until  the  seeds  are  ripe. 

The  hermaphroditism  here  seems  to  arise  by  sexual  substitu- 
tion in  the  stamens,  either  at  the  time  of  origin  of  the  phyllomes, 
or  soon  after. 

A  mainly  pistillate,  scion-propagated,  hybrid  willow  (S. 
lasiandra  Benth.  X  babylonica  L.)  by  staminody  of  the  pistil, 
also  shows  this  sort  of  hermaphroditism  (a,  pi.  20,  fig.  21). 
This  tree  has,  besides,  many  bisexual  flowers  with  normal  organs, 
mostly  pistil  between  two  stamens  (pi.  20,  fig.  20).  This  her- 
maphroditism seems  to  arise  by  reversion. 

This  tree  has  also  purely  male  flowers,  commonly  of  two  or 
three,  but  up  to  five,  stamens,  the  parents  being  pent-  and  tri- 
androus.  Rarely  a  filament  with  double  anther  occurs,  also 
normal  ovary  and  anther  on  one  stalk.  Stamens  are  in  two 
sets:  One,  mostly  in  male  flowers,  in  anthesis  with  the  pistils; 
the  other,  common  in  perfect  flowers,  in  anthesis  later  (pro- 


VOL.  2]          Mott. — Teratology  in  Calif  ornian  Willows.  215 

togyny)  (pi.  20,  fig.  20).  Numerous  variations  occur  of  the 
types  of  flowers  described,  by  different  combinations,  in  one 
flower,  of  the  different  normal  and  abnormal  sex-organs.  Cat- 
kins are  varied ;  so  also  are  branchlets. 

Especially  in  the  perfect  flowers,  there  is  a  tendency  towards 
two  transverse  glands  in  addition  to  the  normal  one  or  two 
median. 

This  tree  has  borne  abnormal  flowers  three  seasons;  most  in 
the  third.  A  neighboring  also  pistillate  scion  tree  of  the  same 
cross  was  apparently  normal  the  first  season,  somewhat  abnormal 
the  third.  A  third  tree,  apparently  normal  the  first  season,  bore 
one  perfect  flower  the  third.  Some  similarly  abnormal,  mainly 
pistillate,  naturally  planted  hybrids  near  by  (with  also  trees 
mainly  staminate)  seem  to  be  the  parent  stock.  Apparently 
scions  perpetuate  the  abnormalities.  Some  of  the  mainly  pistil- 
late naturally  grown  trees  show  also  apparent  reduplication  of 
phyllomes,  e.g.,  in  one  flower,  three  unicarpellary  pistils  (some- 
times hermaphrodite,  sometimes  not) ,  or  two  normal  pistils.  The 
same  trees  bear  also  pistils  in  various  stages  of  division  into  two 
unicarpellary  pistils. 

A  8.  lasiandra,  otherwise  normal,  shows  the  tendency  to  four 
glands  (pi.  19,  fig.  22). 

Viewed  speculatively,  the  abnormalities  seem  to  indicate  that 
the  ancestral  Salix  flower  consisted  of  pistil  and  two  stamens 
with  a  four-part  perianth,  and  that  it  has  become  unisexual  by 
suppression  of  the  organs  of  one  sex  or  the  other;  the  series  of 
monstrous  sex-organs  apparently  shows  that  the  two  carpels  of 
the  normal  pistil  stand  transverse, — etc.  \ 

Of  the  usually  recognized  possible  causes  of  teratological 
phenomena,  hybridization  seems,  here,  to  offer  the  most  likely 
explanation. 


216  University  of  California  Publications.  [BOTANY 

BIBLIOGRAPHY. 

Penzig's  Pflanzen-Teratologie,  Bd.  II,  contains  an  approxi- 
mately complete  list  of  references  to  the  literature  on  teratology 
in  Salix,  up  to  the  time  of  publication  of  this  work,  in  1894,  Of 
the  articles  therein  mentioned,  those  especially  were  searched  for 
in  the  original,  and  are  here  noted,  which  are  accompanied  by 
illustrations.  For  the  period  since  1894,  up  to  March,  1905, 
this  bibliography  pretends  to  be  practically  complete  for  all  arti- 
cles, whether  with  or  without  figures. 

*  Anderson,  C.  L. 

1890.     A  Monoecious  Willow.     Zoe,  I,  p.  41. 
fAubert,  S. 

1894.     In  Arch.  sc.  phys.  nat.,  Geneve  3,  per.,  t.  31,  p.  307. 
Bail. 

1877.     Neuere  Beobachtungen   der  Androgynie  bei   Salicineen.     Schr.   d. 
Phys.-okon.  Gesellscli.  zu  Konigsberg,  18  Jahrg.,  1  Abth.,  pp. 
94-5. 
Brendel,  F. 

1880.     Transformation    of    anthers    into    ovaries.     American    Naturalist, 
XIV,  p.  442. 

Burkill,  I.  H. 

1898.  Changes  in  the  Sex  of  Willows.    Ann.  of  Bot.,  XII,  p.  557. 
Camus,  E.  G. 

1899.  Fleurs   faussement   hermaphrodites   et   anomalies   florales   dans  le 

genre  Salix.     Bull.  Soc.  Bot.  de  Fr.,  XLIV,  pp.  185-91,  pis. 

IV-VI. 
Eichelbaum. 

1888.     Note.     Botan.  CentralU.,  XXXV,  p.  114. 
Hampe. 

1840.  In  Linnaea.     XIV,  pp.  367-8,  followed  by  six  pages  by  publisher, 

Von  Schlechtendal. 
Haring,  J. 

1894.     Abnorme   Katzchenbildungen  bei   Salix   caprea   L.   und  bei   Salix 
cinerea   L.      (Ester.    Bot.    Zeitsch.,    Wien,    XLIV,    pp.    386-7, 
415-18. 
Hartman. 

1841.  Note.    Flora,  XXIV,  p.  199. 

*  Not  referred  to  by  Penzig. 

t  Have  not  seen  the  original  article. 


VOL.  2]          Mott.— Teratology  in  Calif  ornian  Willows.  217 

Hegelmaier,  F. 

1887.  Abnormitaten  einiger  einheimischen  diclinen  Pflanzen.  Jahresb.  d. 
Ver.  f.  Naturk.,  Wiirttemb.,  XLIII,  pp.  307-15,  Taf.  III. 

Heinricher,  E. 

1883.  Eine  Zwitterbliithe  von  Salix  Caprea  L.  Sitz-ber.  d.  K.  ATcad  d. 
Wissensch.,  Wien;  1  Abth.,  Bd.  LXXXVII,  pp.  129-31,  Taf. 
II,  Fig.  10. 

Kirschleger. 

1841.     Note.     Flora,  XXIV,  Literaturberichte,  Nro.  1,  p.  59. 
1845.     Note.     Flora,  XXVIII,  p.  402. 

Leefe,  J.  E. 

1841.  Observations  on  some  curious  metamorphoses  of  the  pistil  in 
Salix  Capraea.  Transact,  of  the  Botan.  Soc.  of  Edinburgh, 
I,  2,  pp.  113-114,  pi.  VI,  figs.  16-21. 

Lowe,  J. 

1856.  On  an  abnormality  in  the  flowers  of  Salix  Andersoniana.  Ann. 
and  Magaz.  of  Nat.  Hist.,  pp.  254-5,  9  figs. 

Masters,  M.  T. 

1869.     Vegetable  Teratology. 

MUller,  H. 

1868.     In  Botan.  Zeitg.,  pp.  843-4.     Tab.  XIV,  B. 

Panek,  J. 

1891.  Weiden  und  Weidenbastarde  aus  der  Umgebung  von  Hohenstadt 
in  Mahren.  (Ester.  Bot.  Zeitsch.,  Wien,  XLIV,  381-5. 

Sauter. 

1837.     Note.     Flora,  XX,  Beibldtter  ler,  Bd.,  p.  40. 

Schimper. 

1829.     In  Flora,  XII,  pp.  422-3. 

Schnitzlein. 

1850.     Note.     Bot.  Zeitg.     VIII,  p.  746,  Taf.  VIII,  Fig.  2-7. 

Shimek,  B. 

1895.  Perfect  flowers  of  Salix  amygdaloides  Ands.  Proc.  Iowa  Ac.  Sc., 
Ill,  pp.  89-90.  2  figs. 

Tausch. 

1833.       Note.     Flora,  XVI,  p.  230. 

Velanovsky,  J. 

1904.  Vergleichende  Studien  iiber  die  Salix-Bliite.  Beihefte  zum  Bot. 
Centralbl.,  XVII,  pp.  123-8.  Taf.  2. 


218  University  of  California  Publications.  [BOTANY 


Von  Schlechtendal.     (See  Hampe.) 

Von  Seemen,  O. 

1886.  Einiges   iiber    abnorme    Bliitenbildungen    bei    den    Weiden.     Ver- 

handlungen  des  Botan.  Ver.  der  Prov.  Brandenburg,  XXVIII, 
pp.  1-14,  Taf.  I.     (48  small  figs.). 

1895.  Abnorme  Bliitenbildungen  bei  einer  Salix  fragilis  L.  (Ester.  Bot. 
Zeitsch.,  Wien,  XLV,  pp.  254-7,  289-95.  Tafn.  XII,  u. 
XIII. 

Wigand,  A. 

1887.  Beitrage   zur    Pflanzen-Teratologie.     Botanische   Hefte,   II,    Mar- 

burg, p.  106  and  pp.  121-22. 


VOL.  2]          Mott.— Teratology  in  Calif  ornian  Willows.  219 


EXPLANATION  OF  PLATES  19  AND  20. 

All  figures  (except  figs.  16-19)  show  the  posterior  side  of  the  flower, 
i.e.,  the  side  which  faces  the  axis  of  the  ament.  Bract,  honey-gland,  and 
lower  part  of  united  filaments  have  been  removed  in  most  cases. 

All  figures  (excepts  figs.  1  A  and  16-19)  circa  10.  Figs.  7  and  22 
somewhat  larger. 


FIG.  1-19  SALIX  LASIOLEPIS  BENTH. 

Figs.  1-15  show  a  series  of  pairs  of  sexual  phyllomes  varying  in  form 
from  two  stamens  to  a  bicarpellary  pistil.  Each  pair  is  constant  in  form, 
from  the  bud,  but  the  series  are  described  as  though  each  member  repre- 
sented a  temporary  stage  in  a  progressive  change  from  stamens  to  pistil 
taking  place  in  a  single  flower.  Fig.  1  from  normal  male  individual; 
figs.  14  and  15  from  normal  female;  figs.  2  and  13  from  abnormal  male. 
Fig.  1.  Normal  male  flower  (from  specimen  No.  5049  in  the  Herbarium 
of  the  University,  collected  at  Willow  Camp,  Marin  Co.,  Calif.) 

a,  anther-lobe,  composed  of  two  pollen-sacs  (microsporangia) 
which  dehisce  by  a  common  slit  above  the  fused  wall  be- 
tween the  pollen-sacs. 

&,  slit  for  escape  of  pollen. 

c,  cylindrical  filaments  united  to  about  the  middle. 

d,  upper  limit  of  normal  union  of  filaments. 

e,  hairy  bract  of  ament,  subtending  the  flower. 

f,  honey-gland  at  base  of  filament. 

Fig.  IA.     Young    diandrous    flower    of    Salix   purpurea   L.,    with    filaments 
united  and  anthers  extrorse. 
a.a.j  anthers. 
Z>,  united  filaments. 

c,  hairy  bract. 

d,  honey  gland. 

Fig.  2.     Beginning  of  transformation. 

a,  grooves  in  filament-forks,  preparatory  to  formation  of 
ovarian  cavity.  Forks  still  free  and  of  normal  length, 
and  entire  flower  otherwise  normal.  The  barren  por- 
tions of  the  forks  are  finally  reduced,  the  connectives 
developing  to  form  the  ovary  while  the  filaments  shorten 
and  complete  their  union  to  form  the  stipe. 


220  University  of  California  Publications.  [BOTANY 

Fig.  3.     Beginning  of  union  of  grooved  filament  forks. 
a,  anterior  margins,  united. 
6,  fc,  posterior  margins,  free. 

c,  c,  filaments,  widened  slightly  medianly  and  with  more  evi- 

dent grooves  than  in  fig.  2. 

d,  beginning  of  ovarian  cavity;  no  ovules  yet. 

e,  connective,  almost  normal. 

f,  f,  anthers,  normal,  but  introrse  and  twisting  to  face  honey- 

gland  as  they  swell  in  maturing. 

Fig.  4.  First  appearance  of  ovules.  (Normal  number  on  a  placenta  is 
six  or  eight,  hence  normal  number  on  a  phyllome  margin  is  three  or 
four,  since  two  margins  unite  to  form  a  placenta). 

a,  a,  ovules,  marginal  and  close  to  base  of  anther-lobes.     That 
these  bodies  are  ovules  beyond  doubt  is  shown  in  the 
enlarged  drawing,  fig.  16. 
&,  b,  developing  marginal  placenta. 

c,  anther,  still  normally  developed. 

d,  stamen,  normal  except  for  groove;  no  ovules. 

Fig.  5.  Ovules  (some  well  developed,  some  abortive)  more  numerous,  and 
on  both  phyllomes.  The  ovules  are  so  closely  associated  with  the 
pollen-sacs  that  some  of  the  ovules  cause  malformations  in  the  pollen- 
sacs,  and  two  ovules  even  protrude  through  the  pollen-sac  tissue. 

a,  ovule,  borne  on  a  connective,  and  protruding  through  the 
wall-tissue  of  pollen-sac   (*),  and  carrying  some  of  the 
adjacent  tissue  outward  with  it  as  it  develops. 
6,  wall-tissue  of  pollen-sac  (i)  adhering  to  base  of  ovule  (a). 

c,  ovule    protruding    through    pollen-sac    tissue    similarly    to 

ovule  (a). 

d,  lobe  in  wall-tissue  of  pollen-sac    (j),  resembling  an  ovule 

in  form  and  probably  caused  by  rudimentary  ovule  on 
connective  beneath.  (Ovule  fails  to  develop).  Iden- 
tity of  tissue  of  lobe  determined  by  its  possession  of  the 
same  peculiar  characters  which  mark  the  normal  pollen- 
sac.  (Fig.  17). 

e,  e,  e,  lobes  in  pollen-sac  wall  similar  to   (d),  but  less  promi- 

nent. 

f,  f,  slits  between  pollen-sacs  at  time  of  dehiscence. 

g,  g,  ovules,  borne  free  from  anthers. 

h,  h,  h,  h,  vascular  strands  of  the  developing  placentae  now 
evident  as  thickenings  on  the  margins  of  the  phyllomes. 
Margins  still  ununited. 


VOL.  2]          Mott.— Teratology  in  Calif ornian  Willows.  221 

Fig.  6.  Broadening  of  the  connectives  to  form  ovary;  and  beginning  of 
style  on  one  phyllome  the  broadening  extends  into  the  still  elongated 
forks.  (The  ovules  are  usually  not  distant  from  the  anthers  as 
here). 

a,  a,  broadened  connectives  and  forks. 
6,  &,  ovules. 

c,  beginning  of  apical  development  of  connective  to  form 
barren  upper  part  of  ovary  and  style  and  stigma. 
Connective  of  the  other  stamen  is  not  yet  so  developed. 

Fig.  7.  Increase  of  carpellary  character  of  one  phyllome,  and  degeneration 
of  its  anther.  Filament  fork  of  this  phyllome  is  now  unelongated, 
and  the  connective  is  shown  to  be  the  chief  seat  of  reproductive 
activity. 

a,  b,  c,  vestiges  of  the  two  pollen-sacs  of  the  left  anther-lobe. 
Eight  anther-lobe  reduced  to  one  small  sac    (d).     Yet 
(a),    (b)   and   (c)   all  contain  normal  pollen, 
a,  left  pollen-sac,  still  well  developed,  but  largely  independent 

of  the  right. 
&,  right  pollen-sac,  not  well  developed. 

c,  lobe  of  (&),  probably,  similar  to  (d),  fig.  5. 

d,  vestigial    pollen-sac    representing    right    anther-lobe.      The 

other  pollen-sac  has  entirely  failed  to  develop.  (Figs. 
17-19  show  this  body  to  be  a  pollen-sac  and  not  an 
ovule,  though  it  is  about  as  minute  as  an  ovule,  and  is 
easily  mistaken  for  one). 

e,  e,  ovules  near  base  of  anther-lobe,  as  before. 

f,  f,  four  ovules,  occupying  former  position  of  pollen-sacs  (this 

seeming  to  indicate  that  connective  and  placental  re- 
gions are  homologous). 

g,  filament-fork  unelongated,  owing  to  decadence  in  develop- 

ment of  pollen-sacs  of  this  phyllome. 

h,  style,  being  apical  development  of  connective  (style  is 
curled  downward;  phyllome  at  this  stage  soon  withers). 

h1,  upper  barren  portion  of  ovary-wall,  this  also  being  de- 
veloped from  the  connective  (corresponds  to  (&), 
fig.  11). 

i,  stigma,  first  clear  definition. 

j,  ruptured  suture. 

~k,  anther,  still  well  developed. 

I,  three  ovules. 

m,  filament  fork,  grooved,  but  still  about  normally  elongated, 

the  male  character  of  the  phyllome  predominating. 
fn,  honey-gland. 

o,  bract. 


222  University  of  California  Publications.  [BOTANY 

Fig.  8.  Ovarian  cavity  now  well  developed,  though  the  swelling  of  the 
enclosed  pollen-sacs  causes  premature  dehiscence,  and  dehiscence  is 
at  the  sutures  instead  of  at  the  midribs.  (The  pollen-sacs  are  usually 
more  degenerate  at  this  stage.  The  ovules  should  be  shown  more 
elongated,  as  possessing  polarity). 

a,  suture  between  phyllomes,  remaining  intact. 

b,  b,  suture  ruptured. 

c,  c,  ovary-wall,    from   later   and    apical    development    of    the 

two  connectives. 

d,  d,  styles    and    stigmas,    not   yet   united    as   in    the    normal 

pistil;    they  are  imperfectly  developed. 

e,  old  crotch  of  filament  forks,  corresponding  to    (d),  fig.   1. 

f,  f,  filament  forks,  both  unelongated. 

g,  g,  pollen-sacs,  dehisced. 

Fig.  9.  A  closed  ovary,  though  still  an  imperfect  one,  has  been  evolved. 
Anthers  practically  lost. 

a,  ovary,  not  yet  tapering  above. 

b,  b,  styles,  malformed  and  not  yet  united,  and  stigmas  not  yet 

clearly  differentiated. 

c,  stipe,  still  abnormally  long. 

d,  d,  phyllomes,    formerly    staminal,    now    predominantly    car- 

pellary.     They  still  show  clearly  that  they  stood  right 
and  left, — transverse. 

e,  posterior  suture  between  phyllomes  (carpels). 

f,  honey-gland. 

g,  bract. 

Fig.  10.  Transition  to  normal  pistil  almost  complete;  ovary  swollen  below 
and  tapering  above,  style  single,  pollen-sacs  have  disappeared  (fig. 
11).  Yet  stipe  is  still  abnormally  long,  and  stigma  has  lobes  right 
and  left,  corresponding  to  phyllomes,  or  carpels. 

a,  swollen  lower  portion  of  ovary  containing  ovules. 

b,  tapering  barren  upper  portion  of   ovary,  to  accommodate 

seed-hairs. 

c,  single  style,  well  defined. 

d,  demarcation  of  style  from  ovary  as  in  fully  normal  pistil 

(d,  fig.  14). 

e,  e,  stigma-lobes,  being  the  ununited  tips  of  the  carpels. 

f,  posterior    suture    (placentiferous    below)    between    carpels 

(corresponds  to  (e),  fig.  9).     Same  on  anterior  wall. 

g,  stipe,  still  abnormally  long. 
h,  bract. 

i,  honey-gland. 


VOL.  2]          Mott. — Teratology  in  Calif ornian  Willoivs.  223 

Fig.  11.  Same  specimen  as  shown  in  fig.  10,  but  with  posterior  half  of 
ovary-wall  removed  to  show  total  lack  of  development  of  pollen-sacs, 
and  typical  position  of  ovules  (median  and  on  lower  half  of  wall)  ; 
the  number  of  ovules  is  normal. 

a,  swollen  lower  portion  of  ovary  containing  ovules. 
&,  tapering  barren  uppe-r  portion   of   ovary,   to   accommodate 
seed-hairs. 

c,  single  style,  well  defined. 

d,  demarcation  of  style  from  ovary  as  in  fully  normal  pistil 

(d,  fig.  14). 

e,  e,  stigma-lobes,  being  the  ununited  tips  of  the  carpels. 

f,  posterior    suture    (placentiferous    below)    between    carpels 

(corresponds  to   (e),  fig.  9).     Same  on  anterior  wall. 

g,  stipe,  still  abnormally  long. 
h,  bract. 

i,  honey-gland. 

j,  ovules  on  anterior  placenta  formed  by  union  of  phyllome 
margins  (ovules  are  marginal  here  as  shown  to  be  the 
case  in  preceding  figures,  as  fig.  7,  (f). 

Fig.  12.  Dehisced  capsule,  showing  completion  of  change  of  two  stamens 
into  a  bicarpellary  pistil.  (Style  and  stigmas  fallen.)  Dehiscence 
is  loculicidal  (at  the  midribs)  for  valves  are  median  while  carpellary 
phyllomes  are  transverse.  In  the  monstrous  forms,  separation  occurs 
between  the  transverse  phyllomes  and  each  remains  intact.  The 
series  shows  that  the  carpels  in  the  normal  flower  in  this  genus  stand 
transverse. 

a,  normal  stipe. 

6,  ovary,  swollen  with  seeds  and  seed-hairs. 

c,  seed-hairs. 

d,  posterior  placenta,  at  suture  between  carpels   (corresponds 

to  /,  fig.  10). 
-*         e,  e,  midribs  of  carpels,  along  upper  part  of  which  dehiscence 

occurs  (dehiscence  loculicidal). 
f,f,  valves,  standing  median  (anterior  and  posterior). 

Fig.  13.  Anterior  view  of  capsule  in  fig.  12,  to  illustrate  typical  position 
of  placenta. 

a,  anterior  valve. 

ft,  placenta  (seeds  removed),  along  middle  of  lower  half  of 
valve. 

c,  lower  limit  of  natural  dehiscence. 

d,  line  of  fracture  in  tearing  away  posterior  valve. 

e,  e,  fragments  of  posterior  valve. 


224  University  of  California  Publications.  [BOTANY 

Fig.  14.     Normal  female  flower  from  typical  tree  of  Salix  lasiolepis  Benth, 
(Strawberry  Creek,  Grounds  of  the  University  of  California.     Fall 
catkin.     September,   1904).     Posterior  view  as  in  preceding  figures. 
a,  swollen  lower  portion  of  ovary,  containing  ovules. 
&,  tapering   barren  upper   portion   of   ovary   to   accommodate 
seed-hairs. 

c,  single  style,  well  defined. 

d,  demarcation  of  style  from  ovary. 

e,  stigma  lobed  transversely  and  only  slightly  lobed  medianly. 
/,  posterior  suture   (placentiferous  below)   between  the  carpel 

appearing  as   a   seam  in   the  wall.     Same   on  anterior 

wall. 

g,  stipe,  short. 
In,  bract. 
i,  honey-gland. 

Fig.  15.     Partly   side-view   of   pistil  in  fig.    14,   to   show   slight   groove   at 
midrib  of  carpel,  along  upper  part  of  which  dehiscence  occurs, 
a,  groove. 
&,  transverse  groove  in  stigma. 

Fig.  16.  Section  of  ovule  (macrosporangium),  optical  longitudinal,  median, 
through  raphe,  to  establish  identity  of  ovules  in  figs.  4-7.  (Oblong 
form  shows  polarity.  Vestigial,  pollen-sacs  and  the  pollen-sac  lobes 
are  more  likely  to  be  isodiametric,  though  in  general  appearance  and 
size  they  are  easily  mistaken  for  ovules). 

a,  nucellus. 

&,  inner  integument. 

c,  outer  integument. 

d,  basal  micropyle  (ovule  anatropus). 

e,  raphe, 

/,  vascular  bundle  in  raphe. 
g,  chalaza. 
h,  funiculus. 
t,  placenta. 

Fig.  17.  Two  cells,  one  from  each  layer  of  wall  of  pollen-sac  in  fig.  19, 
enlarged,  to  show  difference  in  character  of  layers  which  results  in 
dehiscence  when  wall  dries. 

a,  thin-walled  cell   (shown  in  section)   of  outer  layer,  marked 

(6)  in  fig.  19. 
~b,  "fibrous-cell"  of  inner  layer,  marked  (c)  in  fig.  19. 


VOL.  2]          Mott. — Teratology  in  Calif ornian  Willows.  225 

Fig.  18.  Pollen-grain  (microspore)  enlarged  from  vestigial  pollen-sac, 
(d}  fig.  7. 

Fig.  19.  Vestigial  pollen-sac  (microsporangium),  (d)  fig.  7,  enlarged,  to 
show  identifying  characters.  (View  into  the  sac  from  side  attached 
to  connective.) 

a,  line  of  rupture  in  tearing  sac  from  connective. 
Z>,  layer  of  thin-walled  cells  forming  outer  part  of  sac-wall. 
c,  layer    of    so-called   ' '  fibrous-cells ' '    forming  inner   part   of 
sac-wall.     (Unequal  contraction  of   (6)  and  (c)  in  dry- 
ing causes  dehiscence;   (&)  contracts  the  more.) 
dj  pollen-grains  (microspores).     Most  of  the  pollen  has  fallen 
out. 


FIGS.  20,   21.     SALIX  LASIANDEA  BENTH  .X  BABYLONICA  L. 

Fig.  20.  Bisexual  flower,  parts  normally  formed;  predominating  type  of 
flower  on  the  tree.  Glands  often  four,  the  two  not  shown  being 
usually  smaller,  and  transverse. 

a,  pistil  of  two  transverse  carpels. 

6,  suture  between  carpels,  placentiferous  below. 

c,  c,  stamens,    transverse;    only   two,    though    five    normal    for 

S.  lasiandra   (in  this  locality)   in  staminate  flower  and 
three  normal  for  S.  ~babylonica. 

d,  filament,  not  yet  elongated  (flower  protogynous) . 

e,  posterior  honey-gland. 

f,  anterior  honey-gland. 

g,  bract,  subtending  flower. 

Fig.  21.  Bisexual  flower  with  hermaphrodite  organ  in  place  of  pistil ; 
one  of  the  numerous  variations  on  this  tree  of  the  typical  flower, 
fig.  20.  It  shows  one  stage  of  change  of  pistil  to  two  stamens; 
and  verifies  evidence  shown  by  S.  lasiolepis  series  that  carpels  of 
normal  pistil  in  this  genus  stand  transverse. 

a,  monstrous  hermaphrodite  organ. 

b,  1),  phyllomes,  transverse. 

c,  c,  anther-lobes  of  two  pollen-sacs,  each  somewhat  malformed 

and  misplaced  yet  producing  and  discharging  pollen. 

d,  vestigial  anther-lobe. 

e,  two  ovules,  on  anterior  sutural  placenta. 

f,  posterior  suture  of  monstrous  ovary,  prematurely  ruptured 

by  swelling  of  abnormally  enclosed  anthers. 

g,  g,  stamens,  transverse,  with  filaments  not  yet  elongated. 
h,  h,  honey-glands,  median. 


226  University  of  California  Publications.  [BOTANY 

FIG.  22   (PL.  19).     SALIX  LASIANDEA  BENTH. 

Fig.  22.  Staminate  flower  showing  four  glands, — possibly  vestiges  of 
4-part  perianth.  Only  the  posterior  gland  is  commonly  present  in 
this  species;  tree  otherwise  normal.  (The  five  stamens  and  the  bract 
have  been  cut  off.) 

a,  posterior  gland. 

&,  anterior  gland. 

c,  c,  lateral  glands. 

d,  d,  bases  of  five  stamens. 

e,  bract. 


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